39
Iii 128 128~10 Iii ~I~ll w ~I~~I~ lEi
tr~ W Ii
22 Ii ~W 22
iii W W t ~ iii ~ 11 11
1111118 - 1111118
1I1I11~5 1111114 1111116 11111125 1111114 1111116
MICROCOPY RESOLUTION TEST CHART MICROCOPY RESOLUTION TEST CHART NATlON~L 8UR(~lJ OF STANOPDS-1953A NATIONAL BUREAU Of STANDARDS-1963-A
October 1940 Technical Bulletin No 738
DEPolTMIUVTOFAGBItIJLTlJaE
WASRING-TON DCbullmiddotmiddot J o
~
~
InvestigatIons opound the Parasites opound Popillia japonica and Related Scarabaeidae In the Far East poundrOll] 1929 to 1933 inclusivel
By T R GARDXEI2 entomologist and L B PARKER assistant mlomolo7ist Division 0 Fruit Insect Investigations Bureau 0 Entomology and Plant Quarantine t
~ CO~TE~TS
Page Page
Introduction 1 The parasites 0 r~lated ScarahaeicJae 19 The parnsites of Popillia japollica und their Parsi~~ of the lar bull bull 2ll
biologieL middot - -- -- 2 coillclae 20 3 Tiphia aseTicae Allen and Jayne bull 21arasitesorthe adult beetie - shy4 1iphia biuculctla Aile andTachlttloac -- - shy4 1l1rn~s 21
Hamaria inconurufl Walkcr __ CenteteT cinerea Aldrich
5 Tiphia Imrrelli Parker 22 ElliriropTi jarana Town5Cnd 6 Ti phia calalleaerOT Parker 23
Tfpaop clau~eni Aldrich o Ti raquohia homoncllfaris Pllrkcr 23 PTgomiddottrdae 7iphia ioIaainPnrker 21
AdJl8ilia Jladseta Aldrich 7 7iphia lIo1opolila alitlli Hoberts 24 g Tiphiaatoi Parker 25Parnsitesmthe Inrn g TiphialeIII1a Parker bull 26DeXl1d1le S Tiphiu tegitip[uqa Alicn andpUia reniTai Aldrich
Prosena sibirita (FnbriciWi) 10 anes bullbull 26 n Tlphlll sp ralwan bull IScoliidae H Parasites on which no incsti~ations were
Tiphia popilliarom Hohwer Tiphia cernalis Hohwr
Hi carried on during the pltrioc 192ltJ-33 S Tiphia matllTO Allln and Jaynes 17 Re~tn~ ~nlt1 ~hilpin~ oC ~coliidae 29 Tiphia putlieoTu AIIen and Jaynes IS (IoarIng ~co mIne _-- - - -~ - - ~ ~9
Tiphia sp Taiwan 5 IS Shipping cocoons and adult fcmales bull 30 Surnmnr of parasite shipments to the Cnlted
States __ 31 Sumnlnry ~ ___ ~ ~ __ ~ __ _ ~ ~ _~_ ~~ 31 Literature cited __ 35
INTRODUCTlOX
Investigations of the parasit(s of the Jttpanese be(tl (Popillia iaponica ~ewm) were bgul1 in the Far East in thE spring of 1920 and accounts of thpse investigations from 1920 to 1928 inclusive have appeared in previous publications (9 10)3 This bulletin which
I Submitted for publication rar~h 21 190 Transferred to the Di ision of i-oreig-n Pllraltite Introduction Jul- I 193i l ftcr termin~tion f t~e iestieations in Indin In 199 the iunior author was transferred to Japnn
takmg charge of mwsugntlOns In that countrr In wn When the ScOlor lIuthor returned to the (nited tutes bull The authors wish to acknowled~e tho tluable h~lp of R W Burrell who asslst~d ill the work in Japan
and Chosen in the summer Of 1930 and again in 1912 when h wa~ assi~nc(1 to tit work in those countries of O J Haeussler who supcrise1 the investi~nlions in 1931 in conjunction with the parasilC inYe5thmtlons of the oriental fruit moth or K atoassistnnt in Japan sioro l~21 of S Fujii and of H iuliura Duringthecourse of the jnesti~nFions in Jap~n Chosen India fIIl~ Tai~an the wrilces we Jrel1tl~ aided by the variOUS Gowrttmcnt offiCials of those conntrics The liltmiddot [ huwana (hwf IIf the IlIIpenal Quarantine ServIce and his stutl alTered ecry fadlilY for the work in Japan The dirrclor and stnlT Of the Chosln Ag-riclIltural Experiment Station at uigen C hosn huc kinllly prohlccllahoratnry filiIllies c)(h SClSI)tI The director and staff of the fllian Agricultural Explmiddotriment tation at Taihoku Taiwan kincJl [)rmitled tltr usc of tbe station collection for determinauon purpses lh~ work in Indill wus aided tv 1 Bainbrigle Flctcher Imperial Entomologist of that COllntry atlll hy Col [[ [orrison Director oi the Pasteur Institute at Shillong India who prodded lahQrntorycqnipmcnl fllCilities
Italic numbers in parentheses rcfcr to Litcrature Citrd p 35 214374-10-1
2 TECHICAL BULLETIN 738 U S DEPT OF AGRICULTURE
covers the work of the 5-year period from 1929 to 1933 inclusiYe in Japan Chosen India and Taiwan (Formosa) supplements the previous publications and presents additional information relative to the parasites previously listed as well as accounts of other species fOlmd attacking the genus Popillia In addition investigations were mude on the pnrasites of Anomala orientalis middotWaterh Autoserica castanea (Arrow) and Serica peteglina Ohapin6 three additional beetles which have b(come established in the northeastern part of the United States in recent years
The work in Japan Chosen and Taiwnll was cnrriltd on in n mann(l similar to that of pr(oious YNtrs and consisted of collecting and rrarshying parasit(s of Popillia and relat(d SCilrabadctle nnd shipping tll(sP to the enit(d Stutes togltther with further studi(s on the fi(ld abUllshydance and distribution of these pnlasites The investigations in India which cent(red at Shillong Khasi Hills Assam Prooince wpr( continued along lines preyiously d(scrib(d and consistrd of colllcting rearing and shipping parnsites of Popillia At thl pud of 1929 HlC work in that couutrv was terminatld
No iun~stigations~hay( been carried OIl in China since 1926 Wh(ll
the work in that country v-as terminated owing to the unsettlpdpolitical conditions
THE PARASITES OF POPILUA JAPONICA A~D THEIR BIOLOGIES
During the period 1929-33 im-estigations w(re nUld( in JapHIl Ohosen Chime India and lniwan of the parasitps of Popillia be(tl(s inc1ud((1 in tahle 1 the iclc-fL being to introduc( tlHm into tlw Lnit(d Stutes for the control of Popillia iaponica Under normal pound1(ld conclishytions these speci(s are either parasitic on P jnpanica or attack other m(mb(rs of the same genus The deA-iids aI( capablltgt of pnrasitizing other genera in til( scamba(id subfamilies l[do10nthillae and Selishycinae in addition to Popillia
TAlILE I-List of parasites of Popillia investigated in Japan Chosen China India and Taiwan
ParasiteS of- Countrie Hosts
Tb~ adult beNle
T~~Z~~~~nma Ald ___ Jaln_____ bull---------- ____bull___J Popillin japonica E t T i do ( DoTlIop18]ana ons bull__ ____ --------bullbull ------- ----ilAnomllla ~p
lIPPill JapoicaI Popillia q1adriguttllta (F tI I lopillll1 lthintlli~ Frimiddot
i Popiflia ltyaTUa lloP~t Ik J Cb (1 j I d Papi cvprzltuUi J[orlCHamara Inltongrua ~ a ----j apnn osen Inn n la____ Popillia 11IlIian X m I Poprlia m18tulllll Fnirm
i lnoml orklltali Wnwrh ~middottllomala TufocuprfL ~rt)t~I middotldoulllJ ttTwimllCltiatll Waterb
Trophop lt1allscm Ald------- J apan _____ __ _________ _ PopUlia japonira bull I rID mala ruomprrbullbull Pyrgotldne) I
I - 1 JPopillia mpricolibullbull
AdapZla flae eta ld_______ I nd f- - _- ---- _______ 1Pnplila ltjllllla I j popJm mllrleUllnrli Hllpe
In prpiOllS publications (9 10) this species is referred to as P cat1lnopera Hope
I In a previous pllhlilation (10) tbis sptci~s is n(erred to as Sbullbullimili Ltwi
I
3 TXVESTIGATIONS OF PARSITES OF TAPANESE BEETLE
TABLE I-List of parasites of Popilliainvestigated in Japan Chosen China India and Taiwan-Continued
Parasites oi- Conntries HostsI--------------1--------The Jana
Dcxiidae I Popilla quadriguitala
I Popillia utrocoerufa Bales Dexa fenTa Jd Miridiba koreana N and K s A ___________ ChoseD_____________ bull____________ Allomala spp
f IPhvllopertha spp
1 f~WiiJjpmlicaPopillla cllpricolli bull
Proua fmla (F)bull ________ Japan ChOseD I n(lIn __bull______ Popilla sppbull bullbull_ A1omala spp Adoreu5p Seriea sppI I
Scolild~~ 1 I(Popillia cupricolli bullTlpha malura A and L_____ Indla______________________________ Popillia spp
I IjPOJlilllUiapmlicaI Poplllia atrocoerlJlea
Tiphia popiliarora Roh_____ l Japan Chosen China_ bull __bull_____ bull Popillia quadrigllitata bull I Popillia chinibullbull i 1 Popillia (orrnoampalla Arrow
Ti hla pllII d J ____ 1nd middotmiddot_1lPOPiliiPopillaa sppcllpricolli bullbull p rora _ an la__________________________
1 Popillia japoniea Ch I Popillia qundrigultaia Tlph remal ROh_____bull__ Japan Chosen lna__ _____ bull Popillia chineni bullbull
i I Popila sppTiphia sp Taiwan 5________ Taiwan-__________________________l Do
unly th( investigations pertaining to the lifc historils habits field observations and reariIl6 and shipping of these parasites in the Far East are dealt with in this bulletin The methods of handling upon arrival in the Cnited States and the rearing colonization und effectiveshyness of these parasites are dealt with in other publications (18456 11 12 18 14 lij 16 17) emanating from the investigations on the Japanese and ASIatic beetles
PARASITES OF THE ADULT BEETLE
Investigations were carried on with four species of Tachinidae parashysitic on adults of Pop-illia japonica in Japan and one species of Pyrgotidlle parasitic on adu1ts of species of Popillia in India No parllshysites of adult Popillia beetles were found in either Chosen or Taiwan except Hamaria inconflrua which is known to be widely distributed throughout the Asiatic and ~falayan Regions Centeter cinerea in Japan and the pyrgotid Adapsilia_fi(wisela in India have but one generation fL
year and Hle well synchronized with their hosts in time of appeaIilnCe in the field Eutrixopsis javana has but one generation a Y(llr find its life cycle corresponds closely to that of Centeter In Japan Ii inconshygrua and Trophops clauseni have two or more generations a year and are present as adults in the field Oe1 a longer period than alC th(gt Popillia bCCtles thus requiring otlltr sCilrabaeid beetles as ail(rnate hosts These are usually pl(sCnt in suffiCient numb(rs to Cnnble thest tachinids to complete their MC cycles but a limited number or il lnck of alternate hosts in It new (Ilvilonment presents u sCriolls hnndiCup in the estllblis~tTlnt of these two specie
4 TECHNICAL ll1LLETIX 738 L S DEPT OF AGlUCrLTCHE
TACHINIDAE
CENlETER CIXEREA Aldrich
The biology and habits of Ceniefl7 cinerea the m08t cfTectin palllsite of adult beetles of Popillia iaponica in northcrn Japnn I111s bccll fully (leaH with in previous publications (D 10) Thc work during the period 1929-33 consisted in the inYCstigntion of llCV nlen8 in Tnpnn to determine thc relatiye abundancc and dpgrp( of fiplcl pnmsitizntion and in the collection of pnrnsitizpd i)(etles for the shipll1lnt of puparin of C cinerea to the United States
IUYfstigations on the islnnd of Hokknido in northpll1 Tnp~lIl reYCaled that Centeter cinerea was well distributed in pmcticaIly all the localities vhere Popillia beetles were present The nTcmge parasitization in all the localities innstigated was 372 percent during the hltter part of July and early part of August in 1931 No doubt this anrngr would have been considerably higher hnd the obscrvutions lWln made cudirr in the senson since preyious records at Sapporo on tIll Silme island have shown that the parasitization drops of1 very sl1ddenlmiddot Ilfter the first part of August On the island of Honshu just south of Hokkaido the presencp of C cinerea in different sections was distinctI -ariable nne the degree of parasitization considerably lower as Contrnstpd with that found in Hokkaido In no locality wpre more than j) pprcent of the beetles pnrasitizecl The highest Utes of pnrnsitizn tioll wrre found in the hilly and mountainous sections of the northprn and western portion of the island where the win telS arp sCYerl Hnd SHOW is present on the ground throughout the wintCr In thC lowlnnds of the pastern portion of the island where the winters arp mild und little SIlOW
)ccurs beptlc parasitization was found to be considCrubl- lowpr and in the section south of Yokohama no trace of C cinerw WI1S found
1Yhile the distribution of Hamaxia incongwa was blillg innstigatNI in Kyushu in southern Japan in 1032 beCtlps baring eggs sil11ilur to those of Ce7~teter were colIeetNI near Ino and Kobayashi in the mounshytains at the northern end of the island mergrnc( of til( pnmsitls the following year proYed them to be C cinerea This makes thr third island in Japan on whieh 0 cinerea is known to occur and (xUnels the known southfrn range much beyond that previously r((ordpd
Popillia bpctlts beaTing Centeter eggs wCre collrctCd for llI)ring and shipment in ] 929 1930 und 1933 Approximattly 85 p(reent of tllC parasitized beetles produced Centeter- pupa ria During thrsr ylturs 33663 puparia were sbipped to the UnitCd StatCs The 1933 shipshymtnts consisting of 6163 puparia were from brrtlCs (0llpC t ((I in the yicinity of rtsonomiya and Tachikawil T1Enr Tokyo The (entfler ndults occurring in these lo(alities are prpsent in th( fipld tllp lattpr Pfilt of Junp and early in JlIl- or approximntrly 1 month Nlrlipr thun those at Sapporo in northCm Jnpan from whieh all prpyiolls hipnlPnts of pupnrin l11atprial have originat((1 fn virw of this and thl (OITPshysponding COITllatioll with th( timc of rl11prgPlw( of thp hot ( cinerea from this section mny prOY( to be hpttEl sYllchronizpd ill timp ()[ mCrgpnee with POJ1illia japonica in the Cnitcd Statls
The mCtboltis of (ollpcting nnt lpaling WltJ( the samp a thosC rl11shyployltltl in pl(iolls ypars Tn plPparing tilpound pllpnritl [or hipnlCnt great (nre nlS exCrciseu in eliminating as much of the bpptip hody as possiblp
5 IXYESTlGATIOXS OF PARASITES OF JAPAXESE BEETLE
HAMAXIA INCONGRtA Vnlklr
llam(llia inconglua a pmusite of the adults of many species of Scarabutidae is widely distributed throughout the Asiatic and IVlalaYfill Regions find in various tropicn countries In Japan where it occurs as a parasite of Popillia japonica it has two and possibly thrre genErntions a year Bping of crepuscular habitJ it is active mostly during the carlr morning hours and in the cvening Although the lxuct mlthod of parasitization is not known it is prlSUDlN that thc femult dlposits fully deClopN lalYap lither on the host beltle or on the llrnrby foliagr and that tlipse enter the beetle through thc softer portions of t111 body
Although no shipments of IJama11aincongrua have bern made to the Lnitrd StlltlS since 1928 obsllyations on distribution and degree of field pHrnsitizatioIl werc carrid on in Japan during th period 1929-33
During 1930 1931 and H)3~ onl 200000 Popillia beetlls (1(
collected nt 108 localitis lepnsenting a wide range of climatic tlnll enviroIlmeJltal conditions 011 the islnnds of Honshu Sbikoku and Kyushu in lapan These were collected from the middle of Junc to the end of August nod in some instances eol1ections were made at weekly internlls Since it is impossible to distinguish the pnrasitizpltl beetles in the field owing to the lnrvipositing habit of this tachinid all collecttd material was sent to the Yokohama laboratory for obsershyvation Hamaria incongrua was reared from beetles collected at 68 of these localiti(s on til( 3 islnnds Emelgpnee records from this collected mnterial showed that IIamaria was the moll abundant ill the hilly places cOnled with underbrush in the wnrmer lll(ns Field pamsitizatioll wns excP((lingly YftrinbJe in diffplent spdiol1s unci thl genellll 11Wrnge yely low In 110 locality were morl than 20 percent of till beetks pnrusitized and in only 13 localiti(middots WnS pnmsitizatioll a bovp 5 pCrclll t
Pnlnsi tized b(etles WlI( found ill tlll filhl in ((11 tllll lind westcIll Japan thlougholl t tll( season the highest rates of palUsitization occurring during thf latter part of June and early in July find again the latter part of August At Obuse in western Japan 51 percent oi tlle beetles were parllsitized on Jllly 9 1932 Examination of the beetlfs from the same locnlity coll(cted July 25 nnd 29 rCvealed pnrasitizutions of 23 and 32 pNcpnt respctively which incrensed to 143 percrnt of the beetlls rollcted the last week in August Popillia collected at Takao near Tokyo in enstern Japan on July 3 1932 had n parusitizfl tioD of 11 ppl(ent which dlopprd to 12 flnd 14 percen t in (ollctions made on J tlly 19 nnll 28 lesppctiVCly and rose flgain to 43 p(lcent ill a lot collected August] 9 The differencps in these palnsitization r(gtcorc18 are no doubt tlue to the two generations of llamaJmiddotia I1ppearing in thl field at tlwse times
In alN1S wlHre adults of Popillin japonicn arp prCs(nt in thC fidel over a period of ~ months or longer llam(uia in(ongrua is ab to develop two glneratiom on this host but in arcas where this dOlS not occur flltcrnatc scarnbacid host bectles a Ie lleCPSS1UT for the pnS(lshyvation of this species
6 TECHNICAL BULLETIN 738 U S DEPT OF AGRICCLTGHE
EUlRIXOPSIS J VANA Townsend
In 1923 nnd 1928 a few adults of Eutrixopsis javana were reared at the Riverton laboratory from shipmeuts of beetles parasitized by Oenteiel cinerea forwarded from Sapporo Japan in 1922 and 1927
Although several attempts haYc been made to obtain information on this tachinid in Japan nothing relative to its biology or life history has been found A cardul examination of beetles bearing tllcbinid eggs collected at Snpporo in 1928 revealed that all tlll pupuria conshytained within the dead parasitized beetles wen those of (enteter ~10re than 100000 beetles free from external tacbinid eggs were also examined but not a single Eutrixopsis puparium vas found In 1930 approximately 10000 PopilliCL weI( (oUecteci at 9 otl(r localities on Rokkaido Island ~Jore than half of the beetles bore tachinid eggs and the remainder were free from extemal eggs Luter examination of both pnrnsitizpd and unpuIfisitized beetles revealNl nothing but Centeier puparin
Apparently Eutrilopsis at times parasitizes Popillia in the absence of some other host and may be consici(led of little importance as a fnctor in the control of P j(Lponica in Inpnn
TnOlHOPS CLA lHEN Aldrich
Trophops cla1l8eni was first discoerNl in 1928 whln ~OOO Popillia iaponica brrtlps colllClrd at Tokorozawn neaL Tokyo Japan late in June rcvpaled t parasitization of 12 pCLeent Blrths (ollrcted at this arCH 1 wcek latlL hoe-el did not eontain n single individual
In 1930 while the distribution of and fi(lcl pnrnsitizltioll by (enieter cinerea find lIamaria iIlCOnflla WeIe being inv(stigatNl in XagHno prefedurr westprn Japan Troptops wns again pn(ollutrred in two locnliti(s In 1931 it wus found in sCnrnl ndditiollllllolulitips in the sumc prefpcture The highest pnlHsitiznLion WitS found at Obuse middotwherc collections of Popillia in 19~2 yielded 81 nnd i) pereeut on July 9 Hnd ] 3 rrspecti vlly The field pnrnsitization of hcptllS collected in all loealities however uytrngpd only 14 lwrccnt in that year Of seyrrnl Scarabueidae present in the fi(middotld in these localities Anomala lujocuplea appears to bc the only other beetle a(cpptrd by tlus tachinid as a host Although Popiliia heetles collpded at Toko1ozawa latp in Junc in 1928 we1P found to contnin Trophops laryae no pnrnsitiz((l beetl(s wcre found in NngnTlo prei(ctule brforc July 5 Aftel this date the degne of fipld pnrHsitizntion increnscd until the middle of the month aftrl which it grndllnlly dePlined After the first wepk in August no pnrnsitizcdPopillia could bp found although A rllocuprea coll(cted late in Allgust wpre found to rontain larvue of Trophops Colkctions of berthmiddots made in thr middle of July showed the sex ratio of the attacked brctles to be 61=3 pcrcent mnles nnd 387 perc(nl females
Votes on the L1je llistMy of Trophops claU~flti
Thp tachinid Hy Trophops clauReni is npparCntly l10eturnnl in its habits for it has never he(n obs(ry(d in Ih( filld PlrHsitizcd heetles are uSllnlly found in rather hilly pinelS with dtl1s( undelshygrowth indicating fidel habits similar to those of IIamaxia Hppellteel
7 I~TESTIGATIOXS OF PARASITES OF rAPAXESE BEETLE
examinations of the beetles containing the 1aryue of Trophops showed no traces of external rggs so it is nssuinrd that the femnle deposits living larvae but the exuct mllnnel in which it attacks its host or the manner of penetrution by the 1urva into tll( host heet1( has n(ycr been determined The clrath of the bretle occurs when the purasitc larva has reached the eurly purt of th( thin1 instal TIl( period of huva1 clevrlopmcnt in the host body is similar to that of Hamaria incongrua the ttvemgr time from roHection of the b((tles to pupation of the parasitE being 9 clays nnd(l nOlmnl t(mp(Lll tllles pr(yuiling during midsummer In July nnd August the pupnl stagr in Popiliia COWlS a prriod of 15 to 17 clnys Howrycr in Anomala r1fOC1lprea which werE collected in the latter part of August the pupal peliod was 28 clnys
The rxuct llumb(l of generations of Trophops in (8t(111 Jnpull is not known although there arr at l(ust two a year the first on Popillia und the folloing onr or morf on aailable Popillia nne Anomala lujocuprea In 1982 Tlophops en1llgrd dnring th( rnriy part of August from Popillia b(etl(s that had bern collected in the middle of July und pound1 71tfocllprea colledCd in the latter part of August were also found to contain lnrnlc of this parasite
The stntns of this tnchinid ns a pnmsite of Popillia japonica is son1lwhat similnr to that of [Jamajia incongnw although its known distribution is morE limitrd BrcHusr of its low rnJe of field pamsitishyzntion ns ohs(lwd in its natin habitat in Japan nnd its two or more gruelUtions a yrnr conring it prriod of ndult nctiity of 2 months or long(l the alur of Troplwps clallseni ns a pnrasite of P japonica in the rnitecl Stntes is questionabll
In 1932 1 totnl of 287 pupa ria of TropllOp8 clauseni erl shipPNI to the rnitNI StntrR from which only 16 adults (merged in the spring of 1933
PYHGOTlDAE
ADAPILIA FIA VIfltETA Aldrich
A full account of tIl( lif( history nnd genlral hnbits of Adapsilia jialiwta a parasite of adult Popillia blltlrs in Inclin has heen given in a previous publication (10) In Indin this pyrgotid Il~middot is restricted to the hilly sections that nre eoyered with wieldy Spltc(eI plnntings of pine trees and fnirly dll1sr undergrowth of brush nt an (elltion of about )000 flet It has only ol1r glnlration It year the ndults apppuring in thefilld about thl middll of July Its normal host in India is Popilia cllpricolli8 although P tyanea nnt P macclellandi arl occl1siol1itlly pnrllsitizrcl In parnsitizing th( bp(ll( lhr [(mnle ildapnmiddotZia which is diurnal rrmnins quirs((llt on th( foliag( nfar the feeding br(tlr until thr brrtlr tnkrs flight whr[rupon till fI~ immedishyately pOllnc(s upon thr b(rtlr from nhoH It tlwn quickly insClts its egg through [1 long rrlntiwly slend(r and shnrp ovipositor into the mid-dorsal l(gion of the b((tlr nhdom(n Il(nr the junctllle with thl thorax
Thr work in Indin in H)29 was rrvotc1rl to col1((ting hl(ltlr l1EfLl Sbillong lWei sbi pping 1dapsilia pu pnria obtai nrcl from thpsr hl(tlrs to the Cnited Stt1t(S X0 furthr work was don( with this BI)((ies nftrr 1929 owing to th( dis(ontinuntion of thr work in thttt country
Brrt1Es W(Ire collcet(l(l dllling tIl( period from July 25 to August 18
8 TECHXICAL BULLETIN 738 U S DEPT OF GRICFLTCRE
1929 th( majority being brought in from August 1 to 9 th( p(liod when the synchronization of host and parasite is at its best DlIling this time 553885 beetles we1( brought in by the llum(IOtls l1atic collectors FIOIU these 4310 Adapsilia pupalin were obtained giving a parasitization of 078 percent as compared with 053 in 1928
TIlt manuel of renring uncI shipping wus similnr to thnt (mployed in preyious yenls The beetles were confined in hla cardboard boxes of approximately 1 cubic foot capacity Illld to absorb excess moisture the bottoms w(1e lined with fresh moss Apples were used to f(ed the beetles and were l(phnisbed on alternate dllYS After 12 dnys allliye bC(tles which were (lvid(ntly Ullpn1llsitized W(Ie destroy(d nnd the dead olles placed in opm trnys to allow the Atlapsilia which W(lre still ill the Inrvn1 stngr to pupntt Artel pupntion til( UllCOIlshySUlllNl body rontrnts of th( host wrre rellloved to diminnt( decomshyposition gases that might arise durillg shipmrnt to til( detriment of the pllparia The pupnrin v(1e packed betwNm ltl~nS of fresh sphagnum moss in smull vrlltillLted 1ectnnglllnJ tillS and n number of tlllse en tirely surroUlHlpd by 1 3-illClL layer of fllsh mo (1(
pfldwl in illS(Ctploof woodell box(s for shipment One shipnwnt eonsisting of 3ROO pupnrin was SPIlt to tli( rnited
Stnt(s in 1929 This arrind at its dCstinntion Xonmb(r 9 Illvinp bN1l ell rou tp 42 ditys nt it t(mp(rntue of 40deg to 50 0 F
PARASITES OF TilE LAIVA
nEXIlDAE
DEXIA EXTnTIS Aldrich
Delin 1flltrali tilr most rommon of the ChmltHs( dlxiids Wiltshyfirst found Ilt SUiglll ChosllI in 1922 (9) Owing to its (olllplirntld biology no r(Hring work for the shipnwnt of pnmsitiz(d glU bs to t1H lnit(gtd StatN was aWmpted until 1)21) (10) In that Yenr P()pilli(t jnponic(L grubs whirh do not orcur in Chosen (1( eol1letrd filld shipped from nortlH11l Japan to Fukuoka in Kyusitu soutitNll Jnpitn for pnrnsitizlltion by Duia (mnks fonnumiddottkd from Chos) By thr tinlP n suiJici(nt numb(middot of glflYid f(BmTes Wtl( nvailablr Inte i11 JU11r how(yrr most of the grubs w(re too far d(vdop(d for pnrnshysitiztltion purPOR(S find only 850 pnrnsitizld grubs were shipped to til( enitrd States
In 1926 it WfiS d((idpd to (nlTY on rlaring work in 0hos1 utilizing the nutiY( ChOSlll(RC host grubs This provrd to be very sucressful and OWl 20000 grubs were pamsitizrd flnd shippld to titC rnit((l States in 102(3 lind 1927 B(ctlUSC oJ til( dlLngPl iIlYoiwcl in shipping to the rnited Statts Jive grubs othrl thtlll tilos( of POJlillia iapflnica owing to tIl( risk of 111 rscnpl of thrse sp(cils no l(uring w(lIk WIlf
cHrrild on in Chosen in 1928 but 11 mdhod WIIS dlvCloppd whplpby adult fCmale DIJia could be sitipPllaquo LO Yokollilma SII(clssfully and in 1029 rrnring work wns rnrri(d on til(lC the grll bs of P japof ira being IIs((I tiS hosts
Tabl( 2 shows the slllTinl of ndult f(malCs of [Jain middot(lltr(1Ii forshywllrd((l from SlIig(Il Chospn to YokohfilJ11L in 1029 Hnl 1930 til( yciUS that rCaring work WilS el1JTi(d OIl TIle I1dults PI( ronfillNI in thl shipping containl1s which WCI( similm to thos llIlployltd ill transporting Scoliidne npprox11llat(ly 7 days from the timr o( (oll(eshytion to the tim( of removaL
IXVESTIGATIOXS OF PARASITES OF TAPANK~ BEETLE 9
TABLE 2-SuTviLal of adult females of DeJia ventralis shipped from 81tigcn Chosen 10 Yokohama Japa11in 1929 and 1930
Adults Adults surllingYear Generntion shipped shipment
Numb lrumber PercentI
1929_ bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull _ FirsLbullbullbullbullbullbullbullbullbullbullbullbull _bull __ bull __ bullbull_1 063 552 833 1921 __ bullbullbullbullbull ______ bull __ bullbull ____ lhird____ __ 2281 1800 831 1930 __ _bullbullbullbullbullbull __ bull __ First _______ __ ~9() 6S- 68 1910_ _ __ _____ bull_____ __ 1 erond_____ __ __ 2051 14il 720 1930bullbullbull __ bullbullbullbull __ Third __ 2483 2008 809
Total __ __ __ ___ 8368 t 661 ]----()-1 I
It was extremrly difficult to obtain Popillia grubs in any numbers in the --icinity of Yokohamn for rearing work with Deria but this was OYerC0111( by collecting ancl shipping grubs as IHeclNI from Hokshykaido in nortlltrn Japall to Yokohallla n procedure which proyrd to be yery satisfactory and inrxplnsiYe ~0 rearing work was attempted with the s(cond gel1lrntion of DeJia in 1929 oing to the impossibility of s(curing u sufficient number of grubs in IIokkuido during August This was the YNlr of grub scarcity in northern Japall vhen less than 1 percent of the oYCrwintering grubs huying a 2-y(middotnr lifc cyclc (le still in the brynl stage at thn t time of the yeal
The ptlrasitizNl grubs obtain(d tluough parasitization with the first and s(concl g(l1Plntions of DU(L in June nnd August lPspCctiYely wele shipped to thpCnitltC1 Stat(s immpdiatrly aftC1 btinf pnra~itizcd DcYClopment of the parasites took pla(e P11 route nnd the adults emrTgecl sho1tl)- uiter nrriyal at tlHlT destination Grubs containing tbe hibernnting ln1yue of De ia obtuined through pnrasitizatiOIl with the third-g(lHlntion 1Ii(s in 8(ptp111her nnd Octol)(r Pre kPpt OY(1 wintpr at 10kohi1111tl in n wlll-crain(c plot (oY(INI with sod The winttr lllortnlity of these grubs was rnthel high ns only 171 and HL9 pcrcent werc nIl( when dug up [or shipnlPnt in the springs of 1930 lllHl 1931 rlspertiwly
Rwring and Shipping Jfpoundt7wds
The rcuring work with lJJ(l l(fltl(llj1 was accomplisll(d hy using a bnmiddotpding-jnl 111Cthod simillll to that employrd in ]Jle-iolls y(nrs Approximately 50 grubs (Ie placfll in the lonrhalf of a l-qualmiddott glnss jar with soil A fin(-IJ1Psh SCIepn cngp contllining 5 or () lJ(tia f(malp WIlS fittNI into this jur lind rPRtpd on til( soil nnd through this scnPI1 into the soil til( dlpositcd lalTHP pnsNI in palch of the host gruhs 1llP grubs wpl(~ (xtunilHd pneh chty Hlld thORP showing thf pnrnsit( nmiddotpjlntol~ funlwl (1( ((monel nne Tplacpd with frlsh g-rub The flips which Pl( fpd IPg III ll[l with it solution of sunr und watN linmiddotd f[om 3 middot(middotCk tc) J IJ1fll1th in til( cage nnd pnrnitizld flom a5 to )0 pP[cpnt of tlH grub Iach dnv
In pucking the pnrnsitizplt1 gruh 101 shipnwnt iin-lillpd hoxls 1 foot squnrp und Ii inch(s dNp W(IP wwd Each l)()x contnilpd 5 cro-5lction s(ts RPparutd b lin pintp ploviding it COll1pnltllHnt of nhout 1 (lbic inch foJ Pilch of jOIl glubs Fol shipllllllt 2 of thpse hoxps (1( packlrl togtlwl in n (Tong woodpl1 (nsp TIl( soil in which thp gru bs (n pu(k(d had hren thorough Iy fumign tld wi t h (arbon (hsulfid( to (limirmtC any otlw1 inlcl lllntrrial which might htwe
23-13-1----40-2
]0 TECH~ICAL BULLETIN 738 U S DEPT OF AGRICULTURE
been present In table 3 is a summary of shipments of palasitized grubs and the percentages of parasite emcrgenee after arrivul in thl United Stntls
TABLE 3-SlImmary oj shipments oj Popillia japonica grubs l)ara~itizcd willi Deda Ientralis Jrom Japan to the United States 1929-31
Year Contents oC shipment Grubs Parnsites emergingship(led-------1--------------- ---- ______ lVumber lVlLmber I Pentnt
19211-bullbullbullbullbullbullbullbullbullbull_bullbull Grubs pnmsiti1ed with flrstgenerntion Dezia ~ ~500 ~51 1000 1930_bullbullbull __ bull __ bullbullbullbullbull __ Overwintered grubs parnsitized with thirdmiddot 3--
gcnemtionDaiu ~~_~ __ ~~ ___ _ ___ ~ 3000 II 126 1930bullbullbull ___bullbullbullbullbull __bull ___ Grubs pamsitized with lirst-enerntion Detia~~ 8000 1504 18S 1930_____________ ~ _ __ Orubs pllrasitized with ~econd-generntion Drtia 7000 l42J I 203 1930____ __ ~ ~ ~ _ GrUbs pllmsiti1ed with second-genemtion 2500 399 16 (f
Dufu II1931___________ Oerwintered grubs paraSitized with third- 4000 345 8 ( generation Duia I
I 1--------- shyTotaJorneruge~_i- ___ bullbullbullbull __ _ _______ bull ___ ~ ~ ~)(JOOO I 4499middot 155 f i i
Owing to the establishment of Diria lentrali~ in tIl( United Stutes from which materinl for lccolonizlltion was availabll no shipments of this pfllflsitl Wlre made aftlr 1931
PnOSENA SIBII1TA (Fabricins)
Thl dexiid fly Prosena sibi7ita middotliich IS glllclfllly distrihut(d throughout till Asiatic and ~IalaYlln Hegions is an importnnt pilll1shy
sill of PopWia iaponica in Kolth(1l1 Japan It hns olle glllelation a YfIU ami is tllpubl( of palllsitizing and dfwloping Oll numplou sppcils of sealt1bnpid grubs of suitnblr size The life history nnd ililbits of this dexiicl huve been fullv dPidt with in previolls publieashytions (9 10) bull
The shipments from 1923 to H)27 inclusi-e cOl1[-iistN[ of fitldshycolkrtfd grubs of Popillia iap(mit(l whieh showld a getHrnI fitld parflsitization of 10 to 15 percpnt Becaust of till impossibility of detCrting the plesenee of the pUlllsite latnt whieh is nttuchNI to tlH main trnelwal trunk of the host all collpcted glUbs W(I( shipp((L Tllf grubs (re dug in Hoklmido rudy in the spring lind imnHdilltely forwarded to the rnited States arriving thete shortly beforp tll( emlIgenee of the parasites Although this method proed sntisshyfactory the trllnsportntioll costs wPle (xtrCll1(ly high nnd in 1927 it was dfcided to palllsitize till grubs nrtifjeiall)- by 11 mltthod that hnd bCll1 laquo(eloped in 1921 This ms aecomplished by dissecting til( first-stag( larval from tIle oVlsacs of the glUyid femuks and trilllSshy
ferring them to thC host grubs by mcans of a fine blllsh To instlll penetration by the parnsites the grubs with til( smal lnlYllc wIe vlaeed in cclIs of suitable size in wooden bloek fotmiddot it petiot of 24 hours brron being plaecd in thc field whftP tllr oYClwintrlld
Late in the summer of 1928 40596 grubs W(Il artificially PUIllshysitized in 1l0rthCrTJ Japan llnd placed in a prcpnred soil plot w(11 drained coered with sod and surrounded to a depth of 18 inch(s by a fine-mesh screen to pr(vent the gnlbs from wumlcring nwny T11(s( grubs were dug up carly in May 1929 and 19440 were nliv( of whieh 19000 weIpound shipped to the U ni ted Stat(s
In 1929 it was impossibl( to seure sufficient grubs for inoeulatioll work Consequently it was necessary to colleet Prosena femulCs and
IXETrGATIOXS OF PARASITES 01 rAlANFJSFJ DEETLB 11
ship them to the Yokohnma laboTfitory und carryon palasitization work there with available current-year grubs from that vicinity Only 12 p(rcent of the females were alive on arrival at Yokohama Tlus high mortality WitS due to the extremely hot weather prevailing at the time of shipment Out of 364 females received alive 159 contained Inrvae sufficiently developed for this work During the latter part of August 6300 grubs were parasitized and placed in hibernation qunrters As e~perienced with the overwinter~d grubs parasitized with Dexia 1entralis the wintPI mortality was (xceedingly high in this material and only 900 were alive wh(n dug up the followshying spring These with the 19000 shipped in 1929 together rith the 29000 of DeJia tentralis make a total of 48900 grubs containing dexiids thnt were shipped to the United States during the period covcl(d by this bulletin The method of pl1cking tIl( pnrasitized grubs for shipment was the sam( I1S that described for shipment of grubs parnsitized with D ~entrais
No shipments of Prosena sibirita wer( mude aftN 1930 ns this parasite was then established in the Vnit((1 States
SCOLIIDAE
During the course of the investigations from 1920 to 1933 inclusive on the pnrasites of Popillia and related scarabaeid b(etl(s in the Orient a huge number of Scoliidne principally of the genus Tiphia were found and intensively studied ~fost of thtsP species which were consid(recl to be ntw have since been des(ribtd (2 18 19 2021)
In table 4 is presented the 1llown distribution in Tnpllll ChOS(D China Indin nncI Taiwan of all species of Tiphia real(d or shipped sinet the initilltion of the work Of the 29 sp(ei(s list(ci in this tablr 8 nre known to occur ill Japan Chosrn and China 2 in Jupnll nnd ChOS(Il and 1 in ChOS(1l and Chinn No doubt furtlH1 innstigtltions will show a much wider dissemination throughout the Orient Of the species mentioned in this bulletin only liphia bisfculata nnd T tegitiplaga purnsites of Anomala allld occurring in southern Jnpnn have 2 distinct generations a year
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
j
OQ 13
0 0 lts ~
lt ~ a ~ ~ ~ ~
gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
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~ ~ ~
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CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
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October 1940 Technical Bulletin No 738
DEPolTMIUVTOFAGBItIJLTlJaE
WASRING-TON DCbullmiddotmiddot J o
~
~
InvestigatIons opound the Parasites opound Popillia japonica and Related Scarabaeidae In the Far East poundrOll] 1929 to 1933 inclusivel
By T R GARDXEI2 entomologist and L B PARKER assistant mlomolo7ist Division 0 Fruit Insect Investigations Bureau 0 Entomology and Plant Quarantine t
~ CO~TE~TS
Page Page
Introduction 1 The parasites 0 r~lated ScarahaeicJae 19 The parnsites of Popillia japollica und their Parsi~~ of the lar bull bull 2ll
biologieL middot - -- -- 2 coillclae 20 3 Tiphia aseTicae Allen and Jayne bull 21arasitesorthe adult beetie - shy4 1iphia biuculctla Aile andTachlttloac -- - shy4 1l1rn~s 21
Hamaria inconurufl Walkcr __ CenteteT cinerea Aldrich
5 Tiphia Imrrelli Parker 22 ElliriropTi jarana Town5Cnd 6 Ti phia calalleaerOT Parker 23
Tfpaop clau~eni Aldrich o Ti raquohia homoncllfaris Pllrkcr 23 PTgomiddottrdae 7iphia ioIaainPnrker 21
AdJl8ilia Jladseta Aldrich 7 7iphia lIo1opolila alitlli Hoberts 24 g Tiphiaatoi Parker 25Parnsitesmthe Inrn g TiphialeIII1a Parker bull 26DeXl1d1le S Tiphiu tegitip[uqa Alicn andpUia reniTai Aldrich
Prosena sibirita (FnbriciWi) 10 anes bullbull 26 n Tlphlll sp ralwan bull IScoliidae H Parasites on which no incsti~ations were
Tiphia popilliarom Hohwer Tiphia cernalis Hohwr
Hi carried on during the pltrioc 192ltJ-33 S Tiphia matllTO Allln and Jaynes 17 Re~tn~ ~nlt1 ~hilpin~ oC ~coliidae 29 Tiphia putlieoTu AIIen and Jaynes IS (IoarIng ~co mIne _-- - - -~ - - ~ ~9
Tiphia sp Taiwan 5 IS Shipping cocoons and adult fcmales bull 30 Surnmnr of parasite shipments to the Cnlted
States __ 31 Sumnlnry ~ ___ ~ ~ __ ~ __ _ ~ ~ _~_ ~~ 31 Literature cited __ 35
INTRODUCTlOX
Investigations of the parasit(s of the Jttpanese be(tl (Popillia iaponica ~ewm) were bgul1 in the Far East in thE spring of 1920 and accounts of thpse investigations from 1920 to 1928 inclusive have appeared in previous publications (9 10)3 This bulletin which
I Submitted for publication rar~h 21 190 Transferred to the Di ision of i-oreig-n Pllraltite Introduction Jul- I 193i l ftcr termin~tion f t~e iestieations in Indin In 199 the iunior author was transferred to Japnn
takmg charge of mwsugntlOns In that countrr In wn When the ScOlor lIuthor returned to the (nited tutes bull The authors wish to acknowled~e tho tluable h~lp of R W Burrell who asslst~d ill the work in Japan
and Chosen in the summer Of 1930 and again in 1912 when h wa~ assi~nc(1 to tit work in those countries of O J Haeussler who supcrise1 the investi~nlions in 1931 in conjunction with the parasilC inYe5thmtlons of the oriental fruit moth or K atoassistnnt in Japan sioro l~21 of S Fujii and of H iuliura Duringthecourse of the jnesti~nFions in Jap~n Chosen India fIIl~ Tai~an the wrilces we Jrel1tl~ aided by the variOUS Gowrttmcnt offiCials of those conntrics The liltmiddot [ huwana (hwf IIf the IlIIpenal Quarantine ServIce and his stutl alTered ecry fadlilY for the work in Japan The dirrclor and stnlT Of the Chosln Ag-riclIltural Experiment Station at uigen C hosn huc kinllly prohlccllahoratnry filiIllies c)(h SClSI)tI The director and staff of the fllian Agricultural Explmiddotriment tation at Taihoku Taiwan kincJl [)rmitled tltr usc of tbe station collection for determinauon purpses lh~ work in Indill wus aided tv 1 Bainbrigle Flctcher Imperial Entomologist of that COllntry atlll hy Col [[ [orrison Director oi the Pasteur Institute at Shillong India who prodded lahQrntorycqnipmcnl fllCilities
Italic numbers in parentheses rcfcr to Litcrature Citrd p 35 214374-10-1
2 TECHICAL BULLETIN 738 U S DEPT OF AGRICULTURE
covers the work of the 5-year period from 1929 to 1933 inclusiYe in Japan Chosen India and Taiwan (Formosa) supplements the previous publications and presents additional information relative to the parasites previously listed as well as accounts of other species fOlmd attacking the genus Popillia In addition investigations were mude on the pnrasites of Anomala orientalis middotWaterh Autoserica castanea (Arrow) and Serica peteglina Ohapin6 three additional beetles which have b(come established in the northeastern part of the United States in recent years
The work in Japan Chosen and Taiwnll was cnrriltd on in n mann(l similar to that of pr(oious YNtrs and consisted of collecting and rrarshying parasit(s of Popillia and relat(d SCilrabadctle nnd shipping tll(sP to the enit(d Stutes togltther with further studi(s on the fi(ld abUllshydance and distribution of these pnlasites The investigations in India which cent(red at Shillong Khasi Hills Assam Prooince wpr( continued along lines preyiously d(scrib(d and consistrd of colllcting rearing and shipping parnsites of Popillia At thl pud of 1929 HlC work in that couutrv was terminatld
No iun~stigations~hay( been carried OIl in China since 1926 Wh(ll
the work in that country v-as terminated owing to the unsettlpdpolitical conditions
THE PARASITES OF POPILUA JAPONICA A~D THEIR BIOLOGIES
During the period 1929-33 im-estigations w(re nUld( in JapHIl Ohosen Chime India and lniwan of the parasitps of Popillia be(tl(s inc1ud((1 in tahle 1 the iclc-fL being to introduc( tlHm into tlw Lnit(d Stutes for the control of Popillia iaponica Under normal pound1(ld conclishytions these speci(s are either parasitic on P jnpanica or attack other m(mb(rs of the same genus The deA-iids aI( capablltgt of pnrasitizing other genera in til( scamba(id subfamilies l[do10nthillae and Selishycinae in addition to Popillia
TAlILE I-List of parasites of Popillia investigated in Japan Chosen China India and Taiwan
ParasiteS of- Countrie Hosts
Tb~ adult beNle
T~~Z~~~~nma Ald ___ Jaln_____ bull---------- ____bull___J Popillin japonica E t T i do ( DoTlIop18]ana ons bull__ ____ --------bullbull ------- ----ilAnomllla ~p
lIPPill JapoicaI Popillia q1adriguttllta (F tI I lopillll1 lthintlli~ Frimiddot
i Popiflia ltyaTUa lloP~t Ik J Cb (1 j I d Papi cvprzltuUi J[orlCHamara Inltongrua ~ a ----j apnn osen Inn n la____ Popillia 11IlIian X m I Poprlia m18tulllll Fnirm
i lnoml orklltali Wnwrh ~middottllomala TufocuprfL ~rt)t~I middotldoulllJ ttTwimllCltiatll Waterb
Trophop lt1allscm Ald------- J apan _____ __ _________ _ PopUlia japonira bull I rID mala ruomprrbullbull Pyrgotldne) I
I - 1 JPopillia mpricolibullbull
AdapZla flae eta ld_______ I nd f- - _- ---- _______ 1Pnplila ltjllllla I j popJm mllrleUllnrli Hllpe
In prpiOllS publications (9 10) this species is referred to as P cat1lnopera Hope
I In a previous pllhlilation (10) tbis sptci~s is n(erred to as Sbullbullimili Ltwi
I
3 TXVESTIGATIONS OF PARSITES OF TAPANESE BEETLE
TABLE I-List of parasites of Popilliainvestigated in Japan Chosen China India and Taiwan-Continued
Parasites oi- Conntries HostsI--------------1--------The Jana
Dcxiidae I Popilla quadriguitala
I Popillia utrocoerufa Bales Dexa fenTa Jd Miridiba koreana N and K s A ___________ ChoseD_____________ bull____________ Allomala spp
f IPhvllopertha spp
1 f~WiiJjpmlicaPopillla cllpricolli bull
Proua fmla (F)bull ________ Japan ChOseD I n(lIn __bull______ Popilla sppbull bullbull_ A1omala spp Adoreu5p Seriea sppI I
Scolild~~ 1 I(Popillia cupricolli bullTlpha malura A and L_____ Indla______________________________ Popillia spp
I IjPOJlilllUiapmlicaI Poplllia atrocoerlJlea
Tiphia popiliarora Roh_____ l Japan Chosen China_ bull __bull_____ bull Popillia quadrigllitata bull I Popillia chinibullbull i 1 Popillia (orrnoampalla Arrow
Ti hla pllII d J ____ 1nd middotmiddot_1lPOPiliiPopillaa sppcllpricolli bullbull p rora _ an la__________________________
1 Popillia japoniea Ch I Popillia qundrigultaia Tlph remal ROh_____bull__ Japan Chosen lna__ _____ bull Popillia chineni bullbull
i I Popila sppTiphia sp Taiwan 5________ Taiwan-__________________________l Do
unly th( investigations pertaining to the lifc historils habits field observations and reariIl6 and shipping of these parasites in the Far East are dealt with in this bulletin The methods of handling upon arrival in the Cnited States and the rearing colonization und effectiveshyness of these parasites are dealt with in other publications (18456 11 12 18 14 lij 16 17) emanating from the investigations on the Japanese and ASIatic beetles
PARASITES OF THE ADULT BEETLE
Investigations were carried on with four species of Tachinidae parashysitic on adults of Pop-illia japonica in Japan and one species of Pyrgotidlle parasitic on adu1ts of species of Popillia in India No parllshysites of adult Popillia beetles were found in either Chosen or Taiwan except Hamaria inconflrua which is known to be widely distributed throughout the Asiatic and ~falayan Regions Centeter cinerea in Japan and the pyrgotid Adapsilia_fi(wisela in India have but one generation fL
year and Hle well synchronized with their hosts in time of appeaIilnCe in the field Eutrixopsis javana has but one generation a Y(llr find its life cycle corresponds closely to that of Centeter In Japan Ii inconshygrua and Trophops clauseni have two or more generations a year and are present as adults in the field Oe1 a longer period than alC th(gt Popillia bCCtles thus requiring otlltr sCilrabaeid beetles as ail(rnate hosts These are usually pl(sCnt in suffiCient numb(rs to Cnnble thest tachinids to complete their MC cycles but a limited number or il lnck of alternate hosts in It new (Ilvilonment presents u sCriolls hnndiCup in the estllblis~tTlnt of these two specie
4 TECHNICAL ll1LLETIX 738 L S DEPT OF AGlUCrLTCHE
TACHINIDAE
CENlETER CIXEREA Aldrich
The biology and habits of Ceniefl7 cinerea the m08t cfTectin palllsite of adult beetles of Popillia iaponica in northcrn Japnn I111s bccll fully (leaH with in previous publications (D 10) Thc work during the period 1929-33 consisted in the inYCstigntion of llCV nlen8 in Tnpnn to determine thc relatiye abundancc and dpgrp( of fiplcl pnmsitizntion and in the collection of pnrnsitizpd i)(etles for the shipll1lnt of puparin of C cinerea to the United States
IUYfstigations on the islnnd of Hokknido in northpll1 Tnp~lIl reYCaled that Centeter cinerea was well distributed in pmcticaIly all the localities vhere Popillia beetles were present The nTcmge parasitization in all the localities innstigated was 372 percent during the hltter part of July and early part of August in 1931 No doubt this anrngr would have been considerably higher hnd the obscrvutions lWln made cudirr in the senson since preyious records at Sapporo on tIll Silme island have shown that the parasitization drops of1 very sl1ddenlmiddot Ilfter the first part of August On the island of Honshu just south of Hokkaido the presencp of C cinerea in different sections was distinctI -ariable nne the degree of parasitization considerably lower as Contrnstpd with that found in Hokkaido In no locality wpre more than j) pprcent of the beetles pnrasitizecl The highest Utes of pnrnsitizn tioll wrre found in the hilly and mountainous sections of the northprn and western portion of the island where the win telS arp sCYerl Hnd SHOW is present on the ground throughout the wintCr In thC lowlnnds of the pastern portion of the island where the winters arp mild und little SIlOW
)ccurs beptlc parasitization was found to be considCrubl- lowpr and in the section south of Yokohama no trace of C cinerw WI1S found
1Yhile the distribution of Hamaxia incongwa was blillg innstigatNI in Kyushu in southern Japan in 1032 beCtlps baring eggs sil11ilur to those of Ce7~teter were colIeetNI near Ino and Kobayashi in the mounshytains at the northern end of the island mergrnc( of til( pnmsitls the following year proYed them to be C cinerea This makes thr third island in Japan on whieh 0 cinerea is known to occur and (xUnels the known southfrn range much beyond that previously r((ordpd
Popillia bpctlts beaTing Centeter eggs wCre collrctCd for llI)ring and shipment in ] 929 1930 und 1933 Approximattly 85 p(reent of tllC parasitized beetles produced Centeter- pupa ria During thrsr ylturs 33663 puparia were sbipped to the UnitCd StatCs The 1933 shipshymtnts consisting of 6163 puparia were from brrtlCs (0llpC t ((I in the yicinity of rtsonomiya and Tachikawil T1Enr Tokyo The (entfler ndults occurring in these lo(alities are prpsent in th( fipld tllp lattpr Pfilt of Junp and early in JlIl- or approximntrly 1 month Nlrlipr thun those at Sapporo in northCm Jnpan from whieh all prpyiolls hipnlPnts of pupnrin l11atprial have originat((1 fn virw of this and thl (OITPshysponding COITllatioll with th( timc of rl11prgPlw( of thp hot ( cinerea from this section mny prOY( to be hpttEl sYllchronizpd ill timp ()[ mCrgpnee with POJ1illia japonica in the Cnitcd Statls
The mCtboltis of (ollpcting nnt lpaling WltJ( the samp a thosC rl11shyployltltl in pl(iolls ypars Tn plPparing tilpound pllpnritl [or hipnlCnt great (nre nlS exCrciseu in eliminating as much of the bpptip hody as possiblp
5 IXYESTlGATIOXS OF PARASITES OF JAPAXESE BEETLE
HAMAXIA INCONGRtA Vnlklr
llam(llia inconglua a pmusite of the adults of many species of Scarabutidae is widely distributed throughout the Asiatic and IVlalaYfill Regions find in various tropicn countries In Japan where it occurs as a parasite of Popillia japonica it has two and possibly thrre genErntions a year Bping of crepuscular habitJ it is active mostly during the carlr morning hours and in the cvening Although the lxuct mlthod of parasitization is not known it is prlSUDlN that thc femult dlposits fully deClopN lalYap lither on the host beltle or on the llrnrby foliagr and that tlipse enter the beetle through thc softer portions of t111 body
Although no shipments of IJama11aincongrua have bern made to the Lnitrd StlltlS since 1928 obsllyations on distribution and degree of field pHrnsitizatioIl werc carrid on in Japan during th period 1929-33
During 1930 1931 and H)3~ onl 200000 Popillia beetlls (1(
collected nt 108 localitis lepnsenting a wide range of climatic tlnll enviroIlmeJltal conditions 011 the islnnds of Honshu Sbikoku and Kyushu in lapan These were collected from the middle of Junc to the end of August nod in some instances eol1ections were made at weekly internlls Since it is impossible to distinguish the pnrasitizpltl beetles in the field owing to the lnrvipositing habit of this tachinid all collecttd material was sent to the Yokohama laboratory for obsershyvation Hamaria incongrua was reared from beetles collected at 68 of these localiti(s on til( 3 islnnds Emelgpnee records from this collected mnterial showed that IIamaria was the moll abundant ill the hilly places cOnled with underbrush in the wnrmer lll(ns Field pamsitizatioll wns excP((lingly YftrinbJe in diffplent spdiol1s unci thl genellll 11Wrnge yely low In 110 locality were morl than 20 percent of till beetks pnrusitized and in only 13 localiti(middots WnS pnmsitizatioll a bovp 5 pCrclll t
Pnlnsi tized b(etles WlI( found ill tlll filhl in ((11 tllll lind westcIll Japan thlougholl t tll( season the highest rates of palUsitization occurring during thf latter part of June and early in July find again the latter part of August At Obuse in western Japan 51 percent oi tlle beetles were parllsitized on Jllly 9 1932 Examination of the beetlfs from the same locnlity coll(cted July 25 nnd 29 rCvealed pnrasitizutions of 23 and 32 pNcpnt respctively which incrensed to 143 percrnt of the beetlls rollcted the last week in August Popillia collected at Takao near Tokyo in enstern Japan on July 3 1932 had n parusitizfl tioD of 11 ppl(ent which dlopprd to 12 flnd 14 percen t in (ollctions made on J tlly 19 nnll 28 lesppctiVCly and rose flgain to 43 p(lcent ill a lot collected August] 9 The differencps in these palnsitization r(gtcorc18 are no doubt tlue to the two generations of llamaJmiddotia I1ppearing in thl field at tlwse times
In alN1S wlHre adults of Popillin japonicn arp prCs(nt in thC fidel over a period of ~ months or longer llam(uia in(ongrua is ab to develop two glneratiom on this host but in arcas where this dOlS not occur flltcrnatc scarnbacid host bectles a Ie lleCPSS1UT for the pnS(lshyvation of this species
6 TECHNICAL BULLETIN 738 U S DEPT OF AGRICCLTGHE
EUlRIXOPSIS J VANA Townsend
In 1923 nnd 1928 a few adults of Eutrixopsis javana were reared at the Riverton laboratory from shipmeuts of beetles parasitized by Oenteiel cinerea forwarded from Sapporo Japan in 1922 and 1927
Although several attempts haYc been made to obtain information on this tachinid in Japan nothing relative to its biology or life history has been found A cardul examination of beetles bearing tllcbinid eggs collected at Snpporo in 1928 revealed that all tlll pupuria conshytained within the dead parasitized beetles wen those of (enteter ~10re than 100000 beetles free from external tacbinid eggs were also examined but not a single Eutrixopsis puparium vas found In 1930 approximately 10000 PopilliCL weI( (oUecteci at 9 otl(r localities on Rokkaido Island ~Jore than half of the beetles bore tachinid eggs and the remainder were free from extemal eggs Luter examination of both pnrnsitizpd and unpuIfisitized beetles revealNl nothing but Centeier puparin
Apparently Eutrilopsis at times parasitizes Popillia in the absence of some other host and may be consici(led of little importance as a fnctor in the control of P j(Lponica in Inpnn
TnOlHOPS CLA lHEN Aldrich
Trophops cla1l8eni was first discoerNl in 1928 whln ~OOO Popillia iaponica brrtlps colllClrd at Tokorozawn neaL Tokyo Japan late in June rcvpaled t parasitization of 12 pCLeent Blrths (ollrcted at this arCH 1 wcek latlL hoe-el did not eontain n single individual
In 1930 while the distribution of and fi(lcl pnrnsitizltioll by (enieter cinerea find lIamaria iIlCOnflla WeIe being inv(stigatNl in XagHno prefedurr westprn Japan Troptops wns again pn(ollutrred in two locnliti(s In 1931 it wus found in sCnrnl ndditiollllllolulitips in the sumc prefpcture The highest pnlHsitiznLion WitS found at Obuse middotwherc collections of Popillia in 19~2 yielded 81 nnd i) pereeut on July 9 Hnd ] 3 rrspecti vlly The field pnrnsitization of hcptllS collected in all loealities however uytrngpd only 14 lwrccnt in that year Of seyrrnl Scarabueidae present in the fi(middotld in these localities Anomala lujocuplea appears to bc the only other beetle a(cpptrd by tlus tachinid as a host Although Popiliia heetles collpded at Toko1ozawa latp in Junc in 1928 we1P found to contnin Trophops laryae no pnrnsitiz((l beetl(s wcre found in NngnTlo prei(ctule brforc July 5 Aftel this date the degne of fipld pnrHsitizntion increnscd until the middle of the month aftrl which it grndllnlly dePlined After the first wepk in August no pnrnsitizcdPopillia could bp found although A rllocuprea coll(cted late in Allgust wpre found to rontain larvue of Trophops Colkctions of berthmiddots made in thr middle of July showed the sex ratio of the attacked brctles to be 61=3 pcrcent mnles nnd 387 perc(nl females
Votes on the L1je llistMy of Trophops claU~flti
Thp tachinid Hy Trophops clauReni is npparCntly l10eturnnl in its habits for it has never he(n obs(ry(d in Ih( filld PlrHsitizcd heetles are uSllnlly found in rather hilly pinelS with dtl1s( undelshygrowth indicating fidel habits similar to those of IIamaxia Hppellteel
7 I~TESTIGATIOXS OF PARASITES OF rAPAXESE BEETLE
examinations of the beetles containing the 1aryue of Trophops showed no traces of external rggs so it is nssuinrd that the femnle deposits living larvae but the exuct mllnnel in which it attacks its host or the manner of penetrution by the 1urva into tll( host heet1( has n(ycr been determined The clrath of the bretle occurs when the purasitc larva has reached the eurly purt of th( thin1 instal TIl( period of huva1 clevrlopmcnt in the host body is similar to that of Hamaria incongrua the ttvemgr time from roHection of the b((tles to pupation of the parasitE being 9 clays nnd(l nOlmnl t(mp(Lll tllles pr(yuiling during midsummer In July nnd August the pupnl stagr in Popiliia COWlS a prriod of 15 to 17 clnys Howrycr in Anomala r1fOC1lprea which werE collected in the latter part of August the pupal peliod was 28 clnys
The rxuct llumb(l of generations of Trophops in (8t(111 Jnpull is not known although there arr at l(ust two a year the first on Popillia und the folloing onr or morf on aailable Popillia nne Anomala lujocuprea In 1982 Tlophops en1llgrd dnring th( rnriy part of August from Popillia b(etl(s that had bern collected in the middle of July und pound1 71tfocllprea colledCd in the latter part of August were also found to contain lnrnlc of this parasite
The stntns of this tnchinid ns a pnmsite of Popillia japonica is son1lwhat similnr to that of [Jamajia incongnw although its known distribution is morE limitrd BrcHusr of its low rnJe of field pamsitishyzntion ns ohs(lwd in its natin habitat in Japan nnd its two or more gruelUtions a yrnr conring it prriod of ndult nctiity of 2 months or long(l the alur of Troplwps clallseni ns a pnrasite of P japonica in the rnitecl Stntes is questionabll
In 1932 1 totnl of 287 pupa ria of TropllOp8 clauseni erl shipPNI to the rnitNI StntrR from which only 16 adults (merged in the spring of 1933
PYHGOTlDAE
ADAPILIA FIA VIfltETA Aldrich
A full account of tIl( lif( history nnd genlral hnbits of Adapsilia jialiwta a parasite of adult Popillia blltlrs in Inclin has heen given in a previous publication (10) In Indin this pyrgotid Il~middot is restricted to the hilly sections that nre eoyered with wieldy Spltc(eI plnntings of pine trees and fnirly dll1sr undergrowth of brush nt an (elltion of about )000 flet It has only ol1r glnlration It year the ndults apppuring in thefilld about thl middll of July Its normal host in India is Popilia cllpricolli8 although P tyanea nnt P macclellandi arl occl1siol1itlly pnrllsitizrcl In parnsitizing th( bp(ll( lhr [(mnle ildapnmiddotZia which is diurnal rrmnins quirs((llt on th( foliag( nfar the feeding br(tlr until thr brrtlr tnkrs flight whr[rupon till fI~ immedishyately pOllnc(s upon thr b(rtlr from nhoH It tlwn quickly insClts its egg through [1 long rrlntiwly slend(r and shnrp ovipositor into the mid-dorsal l(gion of the b((tlr nhdom(n Il(nr the junctllle with thl thorax
Thr work in Indin in H)29 was rrvotc1rl to col1((ting hl(ltlr l1EfLl Sbillong lWei sbi pping 1dapsilia pu pnria obtai nrcl from thpsr hl(tlrs to the Cnited Stt1t(S X0 furthr work was don( with this BI)((ies nftrr 1929 owing to th( dis(ontinuntion of thr work in thttt country
Brrt1Es W(Ire collcet(l(l dllling tIl( period from July 25 to August 18
8 TECHXICAL BULLETIN 738 U S DEPT OF GRICFLTCRE
1929 th( majority being brought in from August 1 to 9 th( p(liod when the synchronization of host and parasite is at its best DlIling this time 553885 beetles we1( brought in by the llum(IOtls l1atic collectors FIOIU these 4310 Adapsilia pupalin were obtained giving a parasitization of 078 percent as compared with 053 in 1928
TIlt manuel of renring uncI shipping wus similnr to thnt (mployed in preyious yenls The beetles were confined in hla cardboard boxes of approximately 1 cubic foot capacity Illld to absorb excess moisture the bottoms w(1e lined with fresh moss Apples were used to f(ed the beetles and were l(phnisbed on alternate dllYS After 12 dnys allliye bC(tles which were (lvid(ntly Ullpn1llsitized W(Ie destroy(d nnd the dead olles placed in opm trnys to allow the Atlapsilia which W(lre still ill the Inrvn1 stngr to pupntt Artel pupntion til( UllCOIlshySUlllNl body rontrnts of th( host wrre rellloved to diminnt( decomshyposition gases that might arise durillg shipmrnt to til( detriment of the pllparia The pupnrin v(1e packed betwNm ltl~nS of fresh sphagnum moss in smull vrlltillLted 1ectnnglllnJ tillS and n number of tlllse en tirely surroUlHlpd by 1 3-illClL layer of fllsh mo (1(
pfldwl in illS(Ctploof woodell box(s for shipment One shipnwnt eonsisting of 3ROO pupnrin was SPIlt to tli( rnited
Stnt(s in 1929 This arrind at its dCstinntion Xonmb(r 9 Illvinp bN1l ell rou tp 42 ditys nt it t(mp(rntue of 40deg to 50 0 F
PARASITES OF TilE LAIVA
nEXIlDAE
DEXIA EXTnTIS Aldrich
Delin 1flltrali tilr most rommon of the ChmltHs( dlxiids Wiltshyfirst found Ilt SUiglll ChosllI in 1922 (9) Owing to its (olllplirntld biology no r(Hring work for the shipnwnt of pnmsitiz(d glU bs to t1H lnit(gtd StatN was aWmpted until 1)21) (10) In that Yenr P()pilli(t jnponic(L grubs whirh do not orcur in Chosen (1( eol1letrd filld shipped from nortlH11l Japan to Fukuoka in Kyusitu soutitNll Jnpitn for pnrnsitizlltion by Duia (mnks fonnumiddottkd from Chos) By thr tinlP n suiJici(nt numb(middot of glflYid f(BmTes Wtl( nvailablr Inte i11 JU11r how(yrr most of the grubs w(re too far d(vdop(d for pnrnshysitiztltion purPOR(S find only 850 pnrnsitizld grubs were shipped to til( enitrd States
In 1926 it WfiS d((idpd to (nlTY on rlaring work in 0hos1 utilizing the nutiY( ChOSlll(RC host grubs This provrd to be very sucressful and OWl 20000 grubs were pamsitizrd flnd shippld to titC rnit((l States in 102(3 lind 1927 B(ctlUSC oJ til( dlLngPl iIlYoiwcl in shipping to the rnited Statts Jive grubs othrl thtlll tilos( of POJlillia iapflnica owing to tIl( risk of 111 rscnpl of thrse sp(cils no l(uring w(lIk WIlf
cHrrild on in Chosen in 1928 but 11 mdhod WIIS dlvCloppd whplpby adult fCmale DIJia could be sitipPllaquo LO Yokollilma SII(clssfully and in 1029 rrnring work wns rnrri(d on til(lC the grll bs of P japof ira being IIs((I tiS hosts
Tabl( 2 shows the slllTinl of ndult f(malCs of [Jain middot(lltr(1Ii forshywllrd((l from SlIig(Il Chospn to YokohfilJ11L in 1029 Hnl 1930 til( yciUS that rCaring work WilS el1JTi(d OIl TIle I1dults PI( ronfillNI in thl shipping containl1s which WCI( similm to thos llIlployltd ill transporting Scoliidne npprox11llat(ly 7 days from the timr o( (oll(eshytion to the tim( of removaL
IXVESTIGATIOXS OF PARASITES OF TAPANK~ BEETLE 9
TABLE 2-SuTviLal of adult females of DeJia ventralis shipped from 81tigcn Chosen 10 Yokohama Japa11in 1929 and 1930
Adults Adults surllingYear Generntion shipped shipment
Numb lrumber PercentI
1929_ bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull _ FirsLbullbullbullbullbullbullbullbullbullbullbullbull _bull __ bull __ bullbull_1 063 552 833 1921 __ bullbullbullbullbull ______ bull __ bullbull ____ lhird____ __ 2281 1800 831 1930 __ _bullbullbullbullbullbull __ bull __ First _______ __ ~9() 6S- 68 1910_ _ __ _____ bull_____ __ 1 erond_____ __ __ 2051 14il 720 1930bullbullbull __ bullbullbullbull __ Third __ 2483 2008 809
Total __ __ __ ___ 8368 t 661 ]----()-1 I
It was extremrly difficult to obtain Popillia grubs in any numbers in the --icinity of Yokohamn for rearing work with Deria but this was OYerC0111( by collecting ancl shipping grubs as IHeclNI from Hokshykaido in nortlltrn Japall to Yokohallla n procedure which proyrd to be yery satisfactory and inrxplnsiYe ~0 rearing work was attempted with the s(cond gel1lrntion of DeJia in 1929 oing to the impossibility of s(curing u sufficient number of grubs in IIokkuido during August This was the YNlr of grub scarcity in northern Japall vhen less than 1 percent of the oYCrwintering grubs huying a 2-y(middotnr lifc cyclc (le still in the brynl stage at thn t time of the yeal
The ptlrasitizNl grubs obtain(d tluough parasitization with the first and s(concl g(l1Plntions of DU(L in June nnd August lPspCctiYely wele shipped to thpCnitltC1 Stat(s immpdiatrly aftC1 btinf pnra~itizcd DcYClopment of the parasites took pla(e P11 route nnd the adults emrTgecl sho1tl)- uiter nrriyal at tlHlT destination Grubs containing tbe hibernnting ln1yue of De ia obtuined through pnrasitizatiOIl with the third-g(lHlntion 1Ii(s in 8(ptp111her nnd Octol)(r Pre kPpt OY(1 wintpr at 10kohi1111tl in n wlll-crain(c plot (oY(INI with sod The winttr lllortnlity of these grubs was rnthel high ns only 171 and HL9 pcrcent werc nIl( when dug up [or shipnlPnt in the springs of 1930 lllHl 1931 rlspertiwly
Rwring and Shipping Jfpoundt7wds
The rcuring work with lJJ(l l(fltl(llj1 was accomplisll(d hy using a bnmiddotpding-jnl 111Cthod simillll to that employrd in ]Jle-iolls y(nrs Approximately 50 grubs (Ie placfll in the lonrhalf of a l-qualmiddott glnss jar with soil A fin(-IJ1Psh SCIepn cngp contllining 5 or () lJ(tia f(malp WIlS fittNI into this jur lind rPRtpd on til( soil nnd through this scnPI1 into the soil til( dlpositcd lalTHP pnsNI in palch of the host gruhs 1llP grubs wpl(~ (xtunilHd pneh chty Hlld thORP showing thf pnrnsit( nmiddotpjlntol~ funlwl (1( ((monel nne Tplacpd with frlsh g-rub The flips which Pl( fpd IPg III ll[l with it solution of sunr und watN linmiddotd f[om 3 middot(middotCk tc) J IJ1fll1th in til( cage nnd pnrnitizld flom a5 to )0 pP[cpnt of tlH grub Iach dnv
In pucking the pnrnsitizplt1 gruh 101 shipnwnt iin-lillpd hoxls 1 foot squnrp und Ii inch(s dNp W(IP wwd Each l)()x contnilpd 5 cro-5lction s(ts RPparutd b lin pintp ploviding it COll1pnltllHnt of nhout 1 (lbic inch foJ Pilch of jOIl glubs Fol shipllllllt 2 of thpse hoxps (1( packlrl togtlwl in n (Tong woodpl1 (nsp TIl( soil in which thp gru bs (n pu(k(d had hren thorough Iy fumign tld wi t h (arbon (hsulfid( to (limirmtC any otlw1 inlcl lllntrrial which might htwe
23-13-1----40-2
]0 TECH~ICAL BULLETIN 738 U S DEPT OF AGRICULTURE
been present In table 3 is a summary of shipments of palasitized grubs and the percentages of parasite emcrgenee after arrivul in thl United Stntls
TABLE 3-SlImmary oj shipments oj Popillia japonica grubs l)ara~itizcd willi Deda Ientralis Jrom Japan to the United States 1929-31
Year Contents oC shipment Grubs Parnsites emergingship(led-------1--------------- ---- ______ lVumber lVlLmber I Pentnt
19211-bullbullbullbullbullbullbullbullbullbull_bullbull Grubs pnmsiti1ed with flrstgenerntion Dezia ~ ~500 ~51 1000 1930_bullbullbull __ bull __ bullbullbullbullbull __ Overwintered grubs parnsitized with thirdmiddot 3--
gcnemtionDaiu ~~_~ __ ~~ ___ _ ___ ~ 3000 II 126 1930bullbullbull ___bullbullbullbullbull __bull ___ Grubs pamsitized with lirst-enerntion Detia~~ 8000 1504 18S 1930_____________ ~ _ __ Orubs pllrasitized with ~econd-generntion Drtia 7000 l42J I 203 1930____ __ ~ ~ ~ _ GrUbs pllmsiti1ed with second-genemtion 2500 399 16 (f
Dufu II1931___________ Oerwintered grubs paraSitized with third- 4000 345 8 ( generation Duia I
I 1--------- shyTotaJorneruge~_i- ___ bullbullbullbull __ _ _______ bull ___ ~ ~ ~)(JOOO I 4499middot 155 f i i
Owing to the establishment of Diria lentrali~ in tIl( United Stutes from which materinl for lccolonizlltion was availabll no shipments of this pfllflsitl Wlre made aftlr 1931
PnOSENA SIBII1TA (Fabricins)
Thl dexiid fly Prosena sibi7ita middotliich IS glllclfllly distrihut(d throughout till Asiatic and ~IalaYlln Hegions is an importnnt pilll1shy
sill of PopWia iaponica in Kolth(1l1 Japan It hns olle glllelation a YfIU ami is tllpubl( of palllsitizing and dfwloping Oll numplou sppcils of sealt1bnpid grubs of suitnblr size The life history nnd ililbits of this dexiicl huve been fullv dPidt with in previolls publieashytions (9 10) bull
The shipments from 1923 to H)27 inclusi-e cOl1[-iistN[ of fitldshycolkrtfd grubs of Popillia iap(mit(l whieh showld a getHrnI fitld parflsitization of 10 to 15 percpnt Becaust of till impossibility of detCrting the plesenee of the pUlllsite latnt whieh is nttuchNI to tlH main trnelwal trunk of the host all collpcted glUbs W(I( shipp((L Tllf grubs (re dug in Hoklmido rudy in the spring lind imnHdilltely forwarded to the rnited States arriving thete shortly beforp tll( emlIgenee of the parasites Although this method proed sntisshyfactory the trllnsportntioll costs wPle (xtrCll1(ly high nnd in 1927 it was dfcided to palllsitize till grubs nrtifjeiall)- by 11 mltthod that hnd bCll1 laquo(eloped in 1921 This ms aecomplished by dissecting til( first-stag( larval from tIle oVlsacs of the glUyid femuks and trilllSshy
ferring them to thC host grubs by mcans of a fine blllsh To instlll penetration by the parnsites the grubs with til( smal lnlYllc wIe vlaeed in cclIs of suitable size in wooden bloek fotmiddot it petiot of 24 hours brron being plaecd in thc field whftP tllr oYClwintrlld
Late in the summer of 1928 40596 grubs W(Il artificially PUIllshysitized in 1l0rthCrTJ Japan llnd placed in a prcpnred soil plot w(11 drained coered with sod and surrounded to a depth of 18 inch(s by a fine-mesh screen to pr(vent the gnlbs from wumlcring nwny T11(s( grubs were dug up carly in May 1929 and 19440 were nliv( of whieh 19000 weIpound shipped to the U ni ted Stat(s
In 1929 it was impossibl( to seure sufficient grubs for inoeulatioll work Consequently it was necessary to colleet Prosena femulCs and
IXETrGATIOXS OF PARASITES 01 rAlANFJSFJ DEETLB 11
ship them to the Yokohnma laboTfitory und carryon palasitization work there with available current-year grubs from that vicinity Only 12 p(rcent of the females were alive on arrival at Yokohama Tlus high mortality WitS due to the extremely hot weather prevailing at the time of shipment Out of 364 females received alive 159 contained Inrvae sufficiently developed for this work During the latter part of August 6300 grubs were parasitized and placed in hibernation qunrters As e~perienced with the overwinter~d grubs parasitized with Dexia 1entralis the wintPI mortality was (xceedingly high in this material and only 900 were alive wh(n dug up the followshying spring These with the 19000 shipped in 1929 together rith the 29000 of DeJia tentralis make a total of 48900 grubs containing dexiids thnt were shipped to the United States during the period covcl(d by this bulletin The method of pl1cking tIl( pnrasitized grubs for shipment was the sam( I1S that described for shipment of grubs parnsitized with D ~entrais
No shipments of Prosena sibirita wer( mude aftN 1930 ns this parasite was then established in the Vnit((1 States
SCOLIIDAE
During the course of the investigations from 1920 to 1933 inclusive on the pnrasites of Popillia and related scarabaeid b(etl(s in the Orient a huge number of Scoliidne principally of the genus Tiphia were found and intensively studied ~fost of thtsP species which were consid(recl to be ntw have since been des(ribtd (2 18 19 2021)
In table 4 is presented the 1llown distribution in Tnpllll ChOS(D China Indin nncI Taiwan of all species of Tiphia real(d or shipped sinet the initilltion of the work Of the 29 sp(ei(s list(ci in this tablr 8 nre known to occur ill Japan Chosrn and China 2 in Jupnll nnd ChOS(Il and 1 in ChOS(1l and Chinn No doubt furtlH1 innstigtltions will show a much wider dissemination throughout the Orient Of the species mentioned in this bulletin only liphia bisfculata nnd T tegitiplaga purnsites of Anomala allld occurring in southern Jnpnn have 2 distinct generations a year
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
j
OQ 13
0 0 lts ~
lt ~ a ~ ~ ~ ~
gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
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2 TECHICAL BULLETIN 738 U S DEPT OF AGRICULTURE
covers the work of the 5-year period from 1929 to 1933 inclusiYe in Japan Chosen India and Taiwan (Formosa) supplements the previous publications and presents additional information relative to the parasites previously listed as well as accounts of other species fOlmd attacking the genus Popillia In addition investigations were mude on the pnrasites of Anomala orientalis middotWaterh Autoserica castanea (Arrow) and Serica peteglina Ohapin6 three additional beetles which have b(come established in the northeastern part of the United States in recent years
The work in Japan Chosen and Taiwnll was cnrriltd on in n mann(l similar to that of pr(oious YNtrs and consisted of collecting and rrarshying parasit(s of Popillia and relat(d SCilrabadctle nnd shipping tll(sP to the enit(d Stutes togltther with further studi(s on the fi(ld abUllshydance and distribution of these pnlasites The investigations in India which cent(red at Shillong Khasi Hills Assam Prooince wpr( continued along lines preyiously d(scrib(d and consistrd of colllcting rearing and shipping parnsites of Popillia At thl pud of 1929 HlC work in that couutrv was terminatld
No iun~stigations~hay( been carried OIl in China since 1926 Wh(ll
the work in that country v-as terminated owing to the unsettlpdpolitical conditions
THE PARASITES OF POPILUA JAPONICA A~D THEIR BIOLOGIES
During the period 1929-33 im-estigations w(re nUld( in JapHIl Ohosen Chime India and lniwan of the parasitps of Popillia be(tl(s inc1ud((1 in tahle 1 the iclc-fL being to introduc( tlHm into tlw Lnit(d Stutes for the control of Popillia iaponica Under normal pound1(ld conclishytions these speci(s are either parasitic on P jnpanica or attack other m(mb(rs of the same genus The deA-iids aI( capablltgt of pnrasitizing other genera in til( scamba(id subfamilies l[do10nthillae and Selishycinae in addition to Popillia
TAlILE I-List of parasites of Popillia investigated in Japan Chosen China India and Taiwan
ParasiteS of- Countrie Hosts
Tb~ adult beNle
T~~Z~~~~nma Ald ___ Jaln_____ bull---------- ____bull___J Popillin japonica E t T i do ( DoTlIop18]ana ons bull__ ____ --------bullbull ------- ----ilAnomllla ~p
lIPPill JapoicaI Popillia q1adriguttllta (F tI I lopillll1 lthintlli~ Frimiddot
i Popiflia ltyaTUa lloP~t Ik J Cb (1 j I d Papi cvprzltuUi J[orlCHamara Inltongrua ~ a ----j apnn osen Inn n la____ Popillia 11IlIian X m I Poprlia m18tulllll Fnirm
i lnoml orklltali Wnwrh ~middottllomala TufocuprfL ~rt)t~I middotldoulllJ ttTwimllCltiatll Waterb
Trophop lt1allscm Ald------- J apan _____ __ _________ _ PopUlia japonira bull I rID mala ruomprrbullbull Pyrgotldne) I
I - 1 JPopillia mpricolibullbull
AdapZla flae eta ld_______ I nd f- - _- ---- _______ 1Pnplila ltjllllla I j popJm mllrleUllnrli Hllpe
In prpiOllS publications (9 10) this species is referred to as P cat1lnopera Hope
I In a previous pllhlilation (10) tbis sptci~s is n(erred to as Sbullbullimili Ltwi
I
3 TXVESTIGATIONS OF PARSITES OF TAPANESE BEETLE
TABLE I-List of parasites of Popilliainvestigated in Japan Chosen China India and Taiwan-Continued
Parasites oi- Conntries HostsI--------------1--------The Jana
Dcxiidae I Popilla quadriguitala
I Popillia utrocoerufa Bales Dexa fenTa Jd Miridiba koreana N and K s A ___________ ChoseD_____________ bull____________ Allomala spp
f IPhvllopertha spp
1 f~WiiJjpmlicaPopillla cllpricolli bull
Proua fmla (F)bull ________ Japan ChOseD I n(lIn __bull______ Popilla sppbull bullbull_ A1omala spp Adoreu5p Seriea sppI I
Scolild~~ 1 I(Popillia cupricolli bullTlpha malura A and L_____ Indla______________________________ Popillia spp
I IjPOJlilllUiapmlicaI Poplllia atrocoerlJlea
Tiphia popiliarora Roh_____ l Japan Chosen China_ bull __bull_____ bull Popillia quadrigllitata bull I Popillia chinibullbull i 1 Popillia (orrnoampalla Arrow
Ti hla pllII d J ____ 1nd middotmiddot_1lPOPiliiPopillaa sppcllpricolli bullbull p rora _ an la__________________________
1 Popillia japoniea Ch I Popillia qundrigultaia Tlph remal ROh_____bull__ Japan Chosen lna__ _____ bull Popillia chineni bullbull
i I Popila sppTiphia sp Taiwan 5________ Taiwan-__________________________l Do
unly th( investigations pertaining to the lifc historils habits field observations and reariIl6 and shipping of these parasites in the Far East are dealt with in this bulletin The methods of handling upon arrival in the Cnited States and the rearing colonization und effectiveshyness of these parasites are dealt with in other publications (18456 11 12 18 14 lij 16 17) emanating from the investigations on the Japanese and ASIatic beetles
PARASITES OF THE ADULT BEETLE
Investigations were carried on with four species of Tachinidae parashysitic on adults of Pop-illia japonica in Japan and one species of Pyrgotidlle parasitic on adu1ts of species of Popillia in India No parllshysites of adult Popillia beetles were found in either Chosen or Taiwan except Hamaria inconflrua which is known to be widely distributed throughout the Asiatic and ~falayan Regions Centeter cinerea in Japan and the pyrgotid Adapsilia_fi(wisela in India have but one generation fL
year and Hle well synchronized with their hosts in time of appeaIilnCe in the field Eutrixopsis javana has but one generation a Y(llr find its life cycle corresponds closely to that of Centeter In Japan Ii inconshygrua and Trophops clauseni have two or more generations a year and are present as adults in the field Oe1 a longer period than alC th(gt Popillia bCCtles thus requiring otlltr sCilrabaeid beetles as ail(rnate hosts These are usually pl(sCnt in suffiCient numb(rs to Cnnble thest tachinids to complete their MC cycles but a limited number or il lnck of alternate hosts in It new (Ilvilonment presents u sCriolls hnndiCup in the estllblis~tTlnt of these two specie
4 TECHNICAL ll1LLETIX 738 L S DEPT OF AGlUCrLTCHE
TACHINIDAE
CENlETER CIXEREA Aldrich
The biology and habits of Ceniefl7 cinerea the m08t cfTectin palllsite of adult beetles of Popillia iaponica in northcrn Japnn I111s bccll fully (leaH with in previous publications (D 10) Thc work during the period 1929-33 consisted in the inYCstigntion of llCV nlen8 in Tnpnn to determine thc relatiye abundancc and dpgrp( of fiplcl pnmsitizntion and in the collection of pnrnsitizpd i)(etles for the shipll1lnt of puparin of C cinerea to the United States
IUYfstigations on the islnnd of Hokknido in northpll1 Tnp~lIl reYCaled that Centeter cinerea was well distributed in pmcticaIly all the localities vhere Popillia beetles were present The nTcmge parasitization in all the localities innstigated was 372 percent during the hltter part of July and early part of August in 1931 No doubt this anrngr would have been considerably higher hnd the obscrvutions lWln made cudirr in the senson since preyious records at Sapporo on tIll Silme island have shown that the parasitization drops of1 very sl1ddenlmiddot Ilfter the first part of August On the island of Honshu just south of Hokkaido the presencp of C cinerea in different sections was distinctI -ariable nne the degree of parasitization considerably lower as Contrnstpd with that found in Hokkaido In no locality wpre more than j) pprcent of the beetles pnrasitizecl The highest Utes of pnrnsitizn tioll wrre found in the hilly and mountainous sections of the northprn and western portion of the island where the win telS arp sCYerl Hnd SHOW is present on the ground throughout the wintCr In thC lowlnnds of the pastern portion of the island where the winters arp mild und little SIlOW
)ccurs beptlc parasitization was found to be considCrubl- lowpr and in the section south of Yokohama no trace of C cinerw WI1S found
1Yhile the distribution of Hamaxia incongwa was blillg innstigatNI in Kyushu in southern Japan in 1032 beCtlps baring eggs sil11ilur to those of Ce7~teter were colIeetNI near Ino and Kobayashi in the mounshytains at the northern end of the island mergrnc( of til( pnmsitls the following year proYed them to be C cinerea This makes thr third island in Japan on whieh 0 cinerea is known to occur and (xUnels the known southfrn range much beyond that previously r((ordpd
Popillia bpctlts beaTing Centeter eggs wCre collrctCd for llI)ring and shipment in ] 929 1930 und 1933 Approximattly 85 p(reent of tllC parasitized beetles produced Centeter- pupa ria During thrsr ylturs 33663 puparia were sbipped to the UnitCd StatCs The 1933 shipshymtnts consisting of 6163 puparia were from brrtlCs (0llpC t ((I in the yicinity of rtsonomiya and Tachikawil T1Enr Tokyo The (entfler ndults occurring in these lo(alities are prpsent in th( fipld tllp lattpr Pfilt of Junp and early in JlIl- or approximntrly 1 month Nlrlipr thun those at Sapporo in northCm Jnpan from whieh all prpyiolls hipnlPnts of pupnrin l11atprial have originat((1 fn virw of this and thl (OITPshysponding COITllatioll with th( timc of rl11prgPlw( of thp hot ( cinerea from this section mny prOY( to be hpttEl sYllchronizpd ill timp ()[ mCrgpnee with POJ1illia japonica in the Cnitcd Statls
The mCtboltis of (ollpcting nnt lpaling WltJ( the samp a thosC rl11shyployltltl in pl(iolls ypars Tn plPparing tilpound pllpnritl [or hipnlCnt great (nre nlS exCrciseu in eliminating as much of the bpptip hody as possiblp
5 IXYESTlGATIOXS OF PARASITES OF JAPAXESE BEETLE
HAMAXIA INCONGRtA Vnlklr
llam(llia inconglua a pmusite of the adults of many species of Scarabutidae is widely distributed throughout the Asiatic and IVlalaYfill Regions find in various tropicn countries In Japan where it occurs as a parasite of Popillia japonica it has two and possibly thrre genErntions a year Bping of crepuscular habitJ it is active mostly during the carlr morning hours and in the cvening Although the lxuct mlthod of parasitization is not known it is prlSUDlN that thc femult dlposits fully deClopN lalYap lither on the host beltle or on the llrnrby foliagr and that tlipse enter the beetle through thc softer portions of t111 body
Although no shipments of IJama11aincongrua have bern made to the Lnitrd StlltlS since 1928 obsllyations on distribution and degree of field pHrnsitizatioIl werc carrid on in Japan during th period 1929-33
During 1930 1931 and H)3~ onl 200000 Popillia beetlls (1(
collected nt 108 localitis lepnsenting a wide range of climatic tlnll enviroIlmeJltal conditions 011 the islnnds of Honshu Sbikoku and Kyushu in lapan These were collected from the middle of Junc to the end of August nod in some instances eol1ections were made at weekly internlls Since it is impossible to distinguish the pnrasitizpltl beetles in the field owing to the lnrvipositing habit of this tachinid all collecttd material was sent to the Yokohama laboratory for obsershyvation Hamaria incongrua was reared from beetles collected at 68 of these localiti(s on til( 3 islnnds Emelgpnee records from this collected mnterial showed that IIamaria was the moll abundant ill the hilly places cOnled with underbrush in the wnrmer lll(ns Field pamsitizatioll wns excP((lingly YftrinbJe in diffplent spdiol1s unci thl genellll 11Wrnge yely low In 110 locality were morl than 20 percent of till beetks pnrusitized and in only 13 localiti(middots WnS pnmsitizatioll a bovp 5 pCrclll t
Pnlnsi tized b(etles WlI( found ill tlll filhl in ((11 tllll lind westcIll Japan thlougholl t tll( season the highest rates of palUsitization occurring during thf latter part of June and early in July find again the latter part of August At Obuse in western Japan 51 percent oi tlle beetles were parllsitized on Jllly 9 1932 Examination of the beetlfs from the same locnlity coll(cted July 25 nnd 29 rCvealed pnrasitizutions of 23 and 32 pNcpnt respctively which incrensed to 143 percrnt of the beetlls rollcted the last week in August Popillia collected at Takao near Tokyo in enstern Japan on July 3 1932 had n parusitizfl tioD of 11 ppl(ent which dlopprd to 12 flnd 14 percen t in (ollctions made on J tlly 19 nnll 28 lesppctiVCly and rose flgain to 43 p(lcent ill a lot collected August] 9 The differencps in these palnsitization r(gtcorc18 are no doubt tlue to the two generations of llamaJmiddotia I1ppearing in thl field at tlwse times
In alN1S wlHre adults of Popillin japonicn arp prCs(nt in thC fidel over a period of ~ months or longer llam(uia in(ongrua is ab to develop two glneratiom on this host but in arcas where this dOlS not occur flltcrnatc scarnbacid host bectles a Ie lleCPSS1UT for the pnS(lshyvation of this species
6 TECHNICAL BULLETIN 738 U S DEPT OF AGRICCLTGHE
EUlRIXOPSIS J VANA Townsend
In 1923 nnd 1928 a few adults of Eutrixopsis javana were reared at the Riverton laboratory from shipmeuts of beetles parasitized by Oenteiel cinerea forwarded from Sapporo Japan in 1922 and 1927
Although several attempts haYc been made to obtain information on this tachinid in Japan nothing relative to its biology or life history has been found A cardul examination of beetles bearing tllcbinid eggs collected at Snpporo in 1928 revealed that all tlll pupuria conshytained within the dead parasitized beetles wen those of (enteter ~10re than 100000 beetles free from external tacbinid eggs were also examined but not a single Eutrixopsis puparium vas found In 1930 approximately 10000 PopilliCL weI( (oUecteci at 9 otl(r localities on Rokkaido Island ~Jore than half of the beetles bore tachinid eggs and the remainder were free from extemal eggs Luter examination of both pnrnsitizpd and unpuIfisitized beetles revealNl nothing but Centeier puparin
Apparently Eutrilopsis at times parasitizes Popillia in the absence of some other host and may be consici(led of little importance as a fnctor in the control of P j(Lponica in Inpnn
TnOlHOPS CLA lHEN Aldrich
Trophops cla1l8eni was first discoerNl in 1928 whln ~OOO Popillia iaponica brrtlps colllClrd at Tokorozawn neaL Tokyo Japan late in June rcvpaled t parasitization of 12 pCLeent Blrths (ollrcted at this arCH 1 wcek latlL hoe-el did not eontain n single individual
In 1930 while the distribution of and fi(lcl pnrnsitizltioll by (enieter cinerea find lIamaria iIlCOnflla WeIe being inv(stigatNl in XagHno prefedurr westprn Japan Troptops wns again pn(ollutrred in two locnliti(s In 1931 it wus found in sCnrnl ndditiollllllolulitips in the sumc prefpcture The highest pnlHsitiznLion WitS found at Obuse middotwherc collections of Popillia in 19~2 yielded 81 nnd i) pereeut on July 9 Hnd ] 3 rrspecti vlly The field pnrnsitization of hcptllS collected in all loealities however uytrngpd only 14 lwrccnt in that year Of seyrrnl Scarabueidae present in the fi(middotld in these localities Anomala lujocuplea appears to bc the only other beetle a(cpptrd by tlus tachinid as a host Although Popiliia heetles collpded at Toko1ozawa latp in Junc in 1928 we1P found to contnin Trophops laryae no pnrnsitiz((l beetl(s wcre found in NngnTlo prei(ctule brforc July 5 Aftel this date the degne of fipld pnrHsitizntion increnscd until the middle of the month aftrl which it grndllnlly dePlined After the first wepk in August no pnrnsitizcdPopillia could bp found although A rllocuprea coll(cted late in Allgust wpre found to rontain larvue of Trophops Colkctions of berthmiddots made in thr middle of July showed the sex ratio of the attacked brctles to be 61=3 pcrcent mnles nnd 387 perc(nl females
Votes on the L1je llistMy of Trophops claU~flti
Thp tachinid Hy Trophops clauReni is npparCntly l10eturnnl in its habits for it has never he(n obs(ry(d in Ih( filld PlrHsitizcd heetles are uSllnlly found in rather hilly pinelS with dtl1s( undelshygrowth indicating fidel habits similar to those of IIamaxia Hppellteel
7 I~TESTIGATIOXS OF PARASITES OF rAPAXESE BEETLE
examinations of the beetles containing the 1aryue of Trophops showed no traces of external rggs so it is nssuinrd that the femnle deposits living larvae but the exuct mllnnel in which it attacks its host or the manner of penetrution by the 1urva into tll( host heet1( has n(ycr been determined The clrath of the bretle occurs when the purasitc larva has reached the eurly purt of th( thin1 instal TIl( period of huva1 clevrlopmcnt in the host body is similar to that of Hamaria incongrua the ttvemgr time from roHection of the b((tles to pupation of the parasitE being 9 clays nnd(l nOlmnl t(mp(Lll tllles pr(yuiling during midsummer In July nnd August the pupnl stagr in Popiliia COWlS a prriod of 15 to 17 clnys Howrycr in Anomala r1fOC1lprea which werE collected in the latter part of August the pupal peliod was 28 clnys
The rxuct llumb(l of generations of Trophops in (8t(111 Jnpull is not known although there arr at l(ust two a year the first on Popillia und the folloing onr or morf on aailable Popillia nne Anomala lujocuprea In 1982 Tlophops en1llgrd dnring th( rnriy part of August from Popillia b(etl(s that had bern collected in the middle of July und pound1 71tfocllprea colledCd in the latter part of August were also found to contain lnrnlc of this parasite
The stntns of this tnchinid ns a pnmsite of Popillia japonica is son1lwhat similnr to that of [Jamajia incongnw although its known distribution is morE limitrd BrcHusr of its low rnJe of field pamsitishyzntion ns ohs(lwd in its natin habitat in Japan nnd its two or more gruelUtions a yrnr conring it prriod of ndult nctiity of 2 months or long(l the alur of Troplwps clallseni ns a pnrasite of P japonica in the rnitecl Stntes is questionabll
In 1932 1 totnl of 287 pupa ria of TropllOp8 clauseni erl shipPNI to the rnitNI StntrR from which only 16 adults (merged in the spring of 1933
PYHGOTlDAE
ADAPILIA FIA VIfltETA Aldrich
A full account of tIl( lif( history nnd genlral hnbits of Adapsilia jialiwta a parasite of adult Popillia blltlrs in Inclin has heen given in a previous publication (10) In Indin this pyrgotid Il~middot is restricted to the hilly sections that nre eoyered with wieldy Spltc(eI plnntings of pine trees and fnirly dll1sr undergrowth of brush nt an (elltion of about )000 flet It has only ol1r glnlration It year the ndults apppuring in thefilld about thl middll of July Its normal host in India is Popilia cllpricolli8 although P tyanea nnt P macclellandi arl occl1siol1itlly pnrllsitizrcl In parnsitizing th( bp(ll( lhr [(mnle ildapnmiddotZia which is diurnal rrmnins quirs((llt on th( foliag( nfar the feeding br(tlr until thr brrtlr tnkrs flight whr[rupon till fI~ immedishyately pOllnc(s upon thr b(rtlr from nhoH It tlwn quickly insClts its egg through [1 long rrlntiwly slend(r and shnrp ovipositor into the mid-dorsal l(gion of the b((tlr nhdom(n Il(nr the junctllle with thl thorax
Thr work in Indin in H)29 was rrvotc1rl to col1((ting hl(ltlr l1EfLl Sbillong lWei sbi pping 1dapsilia pu pnria obtai nrcl from thpsr hl(tlrs to the Cnited Stt1t(S X0 furthr work was don( with this BI)((ies nftrr 1929 owing to th( dis(ontinuntion of thr work in thttt country
Brrt1Es W(Ire collcet(l(l dllling tIl( period from July 25 to August 18
8 TECHXICAL BULLETIN 738 U S DEPT OF GRICFLTCRE
1929 th( majority being brought in from August 1 to 9 th( p(liod when the synchronization of host and parasite is at its best DlIling this time 553885 beetles we1( brought in by the llum(IOtls l1atic collectors FIOIU these 4310 Adapsilia pupalin were obtained giving a parasitization of 078 percent as compared with 053 in 1928
TIlt manuel of renring uncI shipping wus similnr to thnt (mployed in preyious yenls The beetles were confined in hla cardboard boxes of approximately 1 cubic foot capacity Illld to absorb excess moisture the bottoms w(1e lined with fresh moss Apples were used to f(ed the beetles and were l(phnisbed on alternate dllYS After 12 dnys allliye bC(tles which were (lvid(ntly Ullpn1llsitized W(Ie destroy(d nnd the dead olles placed in opm trnys to allow the Atlapsilia which W(lre still ill the Inrvn1 stngr to pupntt Artel pupntion til( UllCOIlshySUlllNl body rontrnts of th( host wrre rellloved to diminnt( decomshyposition gases that might arise durillg shipmrnt to til( detriment of the pllparia The pupnrin v(1e packed betwNm ltl~nS of fresh sphagnum moss in smull vrlltillLted 1ectnnglllnJ tillS and n number of tlllse en tirely surroUlHlpd by 1 3-illClL layer of fllsh mo (1(
pfldwl in illS(Ctploof woodell box(s for shipment One shipnwnt eonsisting of 3ROO pupnrin was SPIlt to tli( rnited
Stnt(s in 1929 This arrind at its dCstinntion Xonmb(r 9 Illvinp bN1l ell rou tp 42 ditys nt it t(mp(rntue of 40deg to 50 0 F
PARASITES OF TilE LAIVA
nEXIlDAE
DEXIA EXTnTIS Aldrich
Delin 1flltrali tilr most rommon of the ChmltHs( dlxiids Wiltshyfirst found Ilt SUiglll ChosllI in 1922 (9) Owing to its (olllplirntld biology no r(Hring work for the shipnwnt of pnmsitiz(d glU bs to t1H lnit(gtd StatN was aWmpted until 1)21) (10) In that Yenr P()pilli(t jnponic(L grubs whirh do not orcur in Chosen (1( eol1letrd filld shipped from nortlH11l Japan to Fukuoka in Kyusitu soutitNll Jnpitn for pnrnsitizlltion by Duia (mnks fonnumiddottkd from Chos) By thr tinlP n suiJici(nt numb(middot of glflYid f(BmTes Wtl( nvailablr Inte i11 JU11r how(yrr most of the grubs w(re too far d(vdop(d for pnrnshysitiztltion purPOR(S find only 850 pnrnsitizld grubs were shipped to til( enitrd States
In 1926 it WfiS d((idpd to (nlTY on rlaring work in 0hos1 utilizing the nutiY( ChOSlll(RC host grubs This provrd to be very sucressful and OWl 20000 grubs were pamsitizrd flnd shippld to titC rnit((l States in 102(3 lind 1927 B(ctlUSC oJ til( dlLngPl iIlYoiwcl in shipping to the rnited Statts Jive grubs othrl thtlll tilos( of POJlillia iapflnica owing to tIl( risk of 111 rscnpl of thrse sp(cils no l(uring w(lIk WIlf
cHrrild on in Chosen in 1928 but 11 mdhod WIIS dlvCloppd whplpby adult fCmale DIJia could be sitipPllaquo LO Yokollilma SII(clssfully and in 1029 rrnring work wns rnrri(d on til(lC the grll bs of P japof ira being IIs((I tiS hosts
Tabl( 2 shows the slllTinl of ndult f(malCs of [Jain middot(lltr(1Ii forshywllrd((l from SlIig(Il Chospn to YokohfilJ11L in 1029 Hnl 1930 til( yciUS that rCaring work WilS el1JTi(d OIl TIle I1dults PI( ronfillNI in thl shipping containl1s which WCI( similm to thos llIlployltd ill transporting Scoliidne npprox11llat(ly 7 days from the timr o( (oll(eshytion to the tim( of removaL
IXVESTIGATIOXS OF PARASITES OF TAPANK~ BEETLE 9
TABLE 2-SuTviLal of adult females of DeJia ventralis shipped from 81tigcn Chosen 10 Yokohama Japa11in 1929 and 1930
Adults Adults surllingYear Generntion shipped shipment
Numb lrumber PercentI
1929_ bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull _ FirsLbullbullbullbullbullbullbullbullbullbullbullbull _bull __ bull __ bullbull_1 063 552 833 1921 __ bullbullbullbullbull ______ bull __ bullbull ____ lhird____ __ 2281 1800 831 1930 __ _bullbullbullbullbullbull __ bull __ First _______ __ ~9() 6S- 68 1910_ _ __ _____ bull_____ __ 1 erond_____ __ __ 2051 14il 720 1930bullbullbull __ bullbullbullbull __ Third __ 2483 2008 809
Total __ __ __ ___ 8368 t 661 ]----()-1 I
It was extremrly difficult to obtain Popillia grubs in any numbers in the --icinity of Yokohamn for rearing work with Deria but this was OYerC0111( by collecting ancl shipping grubs as IHeclNI from Hokshykaido in nortlltrn Japall to Yokohallla n procedure which proyrd to be yery satisfactory and inrxplnsiYe ~0 rearing work was attempted with the s(cond gel1lrntion of DeJia in 1929 oing to the impossibility of s(curing u sufficient number of grubs in IIokkuido during August This was the YNlr of grub scarcity in northern Japall vhen less than 1 percent of the oYCrwintering grubs huying a 2-y(middotnr lifc cyclc (le still in the brynl stage at thn t time of the yeal
The ptlrasitizNl grubs obtain(d tluough parasitization with the first and s(concl g(l1Plntions of DU(L in June nnd August lPspCctiYely wele shipped to thpCnitltC1 Stat(s immpdiatrly aftC1 btinf pnra~itizcd DcYClopment of the parasites took pla(e P11 route nnd the adults emrTgecl sho1tl)- uiter nrriyal at tlHlT destination Grubs containing tbe hibernnting ln1yue of De ia obtuined through pnrasitizatiOIl with the third-g(lHlntion 1Ii(s in 8(ptp111her nnd Octol)(r Pre kPpt OY(1 wintpr at 10kohi1111tl in n wlll-crain(c plot (oY(INI with sod The winttr lllortnlity of these grubs was rnthel high ns only 171 and HL9 pcrcent werc nIl( when dug up [or shipnlPnt in the springs of 1930 lllHl 1931 rlspertiwly
Rwring and Shipping Jfpoundt7wds
The rcuring work with lJJ(l l(fltl(llj1 was accomplisll(d hy using a bnmiddotpding-jnl 111Cthod simillll to that employrd in ]Jle-iolls y(nrs Approximately 50 grubs (Ie placfll in the lonrhalf of a l-qualmiddott glnss jar with soil A fin(-IJ1Psh SCIepn cngp contllining 5 or () lJ(tia f(malp WIlS fittNI into this jur lind rPRtpd on til( soil nnd through this scnPI1 into the soil til( dlpositcd lalTHP pnsNI in palch of the host gruhs 1llP grubs wpl(~ (xtunilHd pneh chty Hlld thORP showing thf pnrnsit( nmiddotpjlntol~ funlwl (1( ((monel nne Tplacpd with frlsh g-rub The flips which Pl( fpd IPg III ll[l with it solution of sunr und watN linmiddotd f[om 3 middot(middotCk tc) J IJ1fll1th in til( cage nnd pnrnitizld flom a5 to )0 pP[cpnt of tlH grub Iach dnv
In pucking the pnrnsitizplt1 gruh 101 shipnwnt iin-lillpd hoxls 1 foot squnrp und Ii inch(s dNp W(IP wwd Each l)()x contnilpd 5 cro-5lction s(ts RPparutd b lin pintp ploviding it COll1pnltllHnt of nhout 1 (lbic inch foJ Pilch of jOIl glubs Fol shipllllllt 2 of thpse hoxps (1( packlrl togtlwl in n (Tong woodpl1 (nsp TIl( soil in which thp gru bs (n pu(k(d had hren thorough Iy fumign tld wi t h (arbon (hsulfid( to (limirmtC any otlw1 inlcl lllntrrial which might htwe
23-13-1----40-2
]0 TECH~ICAL BULLETIN 738 U S DEPT OF AGRICULTURE
been present In table 3 is a summary of shipments of palasitized grubs and the percentages of parasite emcrgenee after arrivul in thl United Stntls
TABLE 3-SlImmary oj shipments oj Popillia japonica grubs l)ara~itizcd willi Deda Ientralis Jrom Japan to the United States 1929-31
Year Contents oC shipment Grubs Parnsites emergingship(led-------1--------------- ---- ______ lVumber lVlLmber I Pentnt
19211-bullbullbullbullbullbullbullbullbullbull_bullbull Grubs pnmsiti1ed with flrstgenerntion Dezia ~ ~500 ~51 1000 1930_bullbullbull __ bull __ bullbullbullbullbull __ Overwintered grubs parnsitized with thirdmiddot 3--
gcnemtionDaiu ~~_~ __ ~~ ___ _ ___ ~ 3000 II 126 1930bullbullbull ___bullbullbullbullbull __bull ___ Grubs pamsitized with lirst-enerntion Detia~~ 8000 1504 18S 1930_____________ ~ _ __ Orubs pllrasitized with ~econd-generntion Drtia 7000 l42J I 203 1930____ __ ~ ~ ~ _ GrUbs pllmsiti1ed with second-genemtion 2500 399 16 (f
Dufu II1931___________ Oerwintered grubs paraSitized with third- 4000 345 8 ( generation Duia I
I 1--------- shyTotaJorneruge~_i- ___ bullbullbullbull __ _ _______ bull ___ ~ ~ ~)(JOOO I 4499middot 155 f i i
Owing to the establishment of Diria lentrali~ in tIl( United Stutes from which materinl for lccolonizlltion was availabll no shipments of this pfllflsitl Wlre made aftlr 1931
PnOSENA SIBII1TA (Fabricins)
Thl dexiid fly Prosena sibi7ita middotliich IS glllclfllly distrihut(d throughout till Asiatic and ~IalaYlln Hegions is an importnnt pilll1shy
sill of PopWia iaponica in Kolth(1l1 Japan It hns olle glllelation a YfIU ami is tllpubl( of palllsitizing and dfwloping Oll numplou sppcils of sealt1bnpid grubs of suitnblr size The life history nnd ililbits of this dexiicl huve been fullv dPidt with in previolls publieashytions (9 10) bull
The shipments from 1923 to H)27 inclusi-e cOl1[-iistN[ of fitldshycolkrtfd grubs of Popillia iap(mit(l whieh showld a getHrnI fitld parflsitization of 10 to 15 percpnt Becaust of till impossibility of detCrting the plesenee of the pUlllsite latnt whieh is nttuchNI to tlH main trnelwal trunk of the host all collpcted glUbs W(I( shipp((L Tllf grubs (re dug in Hoklmido rudy in the spring lind imnHdilltely forwarded to the rnited States arriving thete shortly beforp tll( emlIgenee of the parasites Although this method proed sntisshyfactory the trllnsportntioll costs wPle (xtrCll1(ly high nnd in 1927 it was dfcided to palllsitize till grubs nrtifjeiall)- by 11 mltthod that hnd bCll1 laquo(eloped in 1921 This ms aecomplished by dissecting til( first-stag( larval from tIle oVlsacs of the glUyid femuks and trilllSshy
ferring them to thC host grubs by mcans of a fine blllsh To instlll penetration by the parnsites the grubs with til( smal lnlYllc wIe vlaeed in cclIs of suitable size in wooden bloek fotmiddot it petiot of 24 hours brron being plaecd in thc field whftP tllr oYClwintrlld
Late in the summer of 1928 40596 grubs W(Il artificially PUIllshysitized in 1l0rthCrTJ Japan llnd placed in a prcpnred soil plot w(11 drained coered with sod and surrounded to a depth of 18 inch(s by a fine-mesh screen to pr(vent the gnlbs from wumlcring nwny T11(s( grubs were dug up carly in May 1929 and 19440 were nliv( of whieh 19000 weIpound shipped to the U ni ted Stat(s
In 1929 it was impossibl( to seure sufficient grubs for inoeulatioll work Consequently it was necessary to colleet Prosena femulCs and
IXETrGATIOXS OF PARASITES 01 rAlANFJSFJ DEETLB 11
ship them to the Yokohnma laboTfitory und carryon palasitization work there with available current-year grubs from that vicinity Only 12 p(rcent of the females were alive on arrival at Yokohama Tlus high mortality WitS due to the extremely hot weather prevailing at the time of shipment Out of 364 females received alive 159 contained Inrvae sufficiently developed for this work During the latter part of August 6300 grubs were parasitized and placed in hibernation qunrters As e~perienced with the overwinter~d grubs parasitized with Dexia 1entralis the wintPI mortality was (xceedingly high in this material and only 900 were alive wh(n dug up the followshying spring These with the 19000 shipped in 1929 together rith the 29000 of DeJia tentralis make a total of 48900 grubs containing dexiids thnt were shipped to the United States during the period covcl(d by this bulletin The method of pl1cking tIl( pnrasitized grubs for shipment was the sam( I1S that described for shipment of grubs parnsitized with D ~entrais
No shipments of Prosena sibirita wer( mude aftN 1930 ns this parasite was then established in the Vnit((1 States
SCOLIIDAE
During the course of the investigations from 1920 to 1933 inclusive on the pnrasites of Popillia and related scarabaeid b(etl(s in the Orient a huge number of Scoliidne principally of the genus Tiphia were found and intensively studied ~fost of thtsP species which were consid(recl to be ntw have since been des(ribtd (2 18 19 2021)
In table 4 is presented the 1llown distribution in Tnpllll ChOS(D China Indin nncI Taiwan of all species of Tiphia real(d or shipped sinet the initilltion of the work Of the 29 sp(ei(s list(ci in this tablr 8 nre known to occur ill Japan Chosrn and China 2 in Jupnll nnd ChOS(Il and 1 in ChOS(1l and Chinn No doubt furtlH1 innstigtltions will show a much wider dissemination throughout the Orient Of the species mentioned in this bulletin only liphia bisfculata nnd T tegitiplaga purnsites of Anomala allld occurring in southern Jnpnn have 2 distinct generations a year
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
j
OQ 13
0 0 lts ~
lt ~ a ~ ~ ~ ~
gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
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34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
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is
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3 TXVESTIGATIONS OF PARSITES OF TAPANESE BEETLE
TABLE I-List of parasites of Popilliainvestigated in Japan Chosen China India and Taiwan-Continued
Parasites oi- Conntries HostsI--------------1--------The Jana
Dcxiidae I Popilla quadriguitala
I Popillia utrocoerufa Bales Dexa fenTa Jd Miridiba koreana N and K s A ___________ ChoseD_____________ bull____________ Allomala spp
f IPhvllopertha spp
1 f~WiiJjpmlicaPopillla cllpricolli bull
Proua fmla (F)bull ________ Japan ChOseD I n(lIn __bull______ Popilla sppbull bullbull_ A1omala spp Adoreu5p Seriea sppI I
Scolild~~ 1 I(Popillia cupricolli bullTlpha malura A and L_____ Indla______________________________ Popillia spp
I IjPOJlilllUiapmlicaI Poplllia atrocoerlJlea
Tiphia popiliarora Roh_____ l Japan Chosen China_ bull __bull_____ bull Popillia quadrigllitata bull I Popillia chinibullbull i 1 Popillia (orrnoampalla Arrow
Ti hla pllII d J ____ 1nd middotmiddot_1lPOPiliiPopillaa sppcllpricolli bullbull p rora _ an la__________________________
1 Popillia japoniea Ch I Popillia qundrigultaia Tlph remal ROh_____bull__ Japan Chosen lna__ _____ bull Popillia chineni bullbull
i I Popila sppTiphia sp Taiwan 5________ Taiwan-__________________________l Do
unly th( investigations pertaining to the lifc historils habits field observations and reariIl6 and shipping of these parasites in the Far East are dealt with in this bulletin The methods of handling upon arrival in the Cnited States and the rearing colonization und effectiveshyness of these parasites are dealt with in other publications (18456 11 12 18 14 lij 16 17) emanating from the investigations on the Japanese and ASIatic beetles
PARASITES OF THE ADULT BEETLE
Investigations were carried on with four species of Tachinidae parashysitic on adults of Pop-illia japonica in Japan and one species of Pyrgotidlle parasitic on adu1ts of species of Popillia in India No parllshysites of adult Popillia beetles were found in either Chosen or Taiwan except Hamaria inconflrua which is known to be widely distributed throughout the Asiatic and ~falayan Regions Centeter cinerea in Japan and the pyrgotid Adapsilia_fi(wisela in India have but one generation fL
year and Hle well synchronized with their hosts in time of appeaIilnCe in the field Eutrixopsis javana has but one generation a Y(llr find its life cycle corresponds closely to that of Centeter In Japan Ii inconshygrua and Trophops clauseni have two or more generations a year and are present as adults in the field Oe1 a longer period than alC th(gt Popillia bCCtles thus requiring otlltr sCilrabaeid beetles as ail(rnate hosts These are usually pl(sCnt in suffiCient numb(rs to Cnnble thest tachinids to complete their MC cycles but a limited number or il lnck of alternate hosts in It new (Ilvilonment presents u sCriolls hnndiCup in the estllblis~tTlnt of these two specie
4 TECHNICAL ll1LLETIX 738 L S DEPT OF AGlUCrLTCHE
TACHINIDAE
CENlETER CIXEREA Aldrich
The biology and habits of Ceniefl7 cinerea the m08t cfTectin palllsite of adult beetles of Popillia iaponica in northcrn Japnn I111s bccll fully (leaH with in previous publications (D 10) Thc work during the period 1929-33 consisted in the inYCstigntion of llCV nlen8 in Tnpnn to determine thc relatiye abundancc and dpgrp( of fiplcl pnmsitizntion and in the collection of pnrnsitizpd i)(etles for the shipll1lnt of puparin of C cinerea to the United States
IUYfstigations on the islnnd of Hokknido in northpll1 Tnp~lIl reYCaled that Centeter cinerea was well distributed in pmcticaIly all the localities vhere Popillia beetles were present The nTcmge parasitization in all the localities innstigated was 372 percent during the hltter part of July and early part of August in 1931 No doubt this anrngr would have been considerably higher hnd the obscrvutions lWln made cudirr in the senson since preyious records at Sapporo on tIll Silme island have shown that the parasitization drops of1 very sl1ddenlmiddot Ilfter the first part of August On the island of Honshu just south of Hokkaido the presencp of C cinerea in different sections was distinctI -ariable nne the degree of parasitization considerably lower as Contrnstpd with that found in Hokkaido In no locality wpre more than j) pprcent of the beetles pnrasitizecl The highest Utes of pnrnsitizn tioll wrre found in the hilly and mountainous sections of the northprn and western portion of the island where the win telS arp sCYerl Hnd SHOW is present on the ground throughout the wintCr In thC lowlnnds of the pastern portion of the island where the winters arp mild und little SIlOW
)ccurs beptlc parasitization was found to be considCrubl- lowpr and in the section south of Yokohama no trace of C cinerw WI1S found
1Yhile the distribution of Hamaxia incongwa was blillg innstigatNI in Kyushu in southern Japan in 1032 beCtlps baring eggs sil11ilur to those of Ce7~teter were colIeetNI near Ino and Kobayashi in the mounshytains at the northern end of the island mergrnc( of til( pnmsitls the following year proYed them to be C cinerea This makes thr third island in Japan on whieh 0 cinerea is known to occur and (xUnels the known southfrn range much beyond that previously r((ordpd
Popillia bpctlts beaTing Centeter eggs wCre collrctCd for llI)ring and shipment in ] 929 1930 und 1933 Approximattly 85 p(reent of tllC parasitized beetles produced Centeter- pupa ria During thrsr ylturs 33663 puparia were sbipped to the UnitCd StatCs The 1933 shipshymtnts consisting of 6163 puparia were from brrtlCs (0llpC t ((I in the yicinity of rtsonomiya and Tachikawil T1Enr Tokyo The (entfler ndults occurring in these lo(alities are prpsent in th( fipld tllp lattpr Pfilt of Junp and early in JlIl- or approximntrly 1 month Nlrlipr thun those at Sapporo in northCm Jnpan from whieh all prpyiolls hipnlPnts of pupnrin l11atprial have originat((1 fn virw of this and thl (OITPshysponding COITllatioll with th( timc of rl11prgPlw( of thp hot ( cinerea from this section mny prOY( to be hpttEl sYllchronizpd ill timp ()[ mCrgpnee with POJ1illia japonica in the Cnitcd Statls
The mCtboltis of (ollpcting nnt lpaling WltJ( the samp a thosC rl11shyployltltl in pl(iolls ypars Tn plPparing tilpound pllpnritl [or hipnlCnt great (nre nlS exCrciseu in eliminating as much of the bpptip hody as possiblp
5 IXYESTlGATIOXS OF PARASITES OF JAPAXESE BEETLE
HAMAXIA INCONGRtA Vnlklr
llam(llia inconglua a pmusite of the adults of many species of Scarabutidae is widely distributed throughout the Asiatic and IVlalaYfill Regions find in various tropicn countries In Japan where it occurs as a parasite of Popillia japonica it has two and possibly thrre genErntions a year Bping of crepuscular habitJ it is active mostly during the carlr morning hours and in the cvening Although the lxuct mlthod of parasitization is not known it is prlSUDlN that thc femult dlposits fully deClopN lalYap lither on the host beltle or on the llrnrby foliagr and that tlipse enter the beetle through thc softer portions of t111 body
Although no shipments of IJama11aincongrua have bern made to the Lnitrd StlltlS since 1928 obsllyations on distribution and degree of field pHrnsitizatioIl werc carrid on in Japan during th period 1929-33
During 1930 1931 and H)3~ onl 200000 Popillia beetlls (1(
collected nt 108 localitis lepnsenting a wide range of climatic tlnll enviroIlmeJltal conditions 011 the islnnds of Honshu Sbikoku and Kyushu in lapan These were collected from the middle of Junc to the end of August nod in some instances eol1ections were made at weekly internlls Since it is impossible to distinguish the pnrasitizpltl beetles in the field owing to the lnrvipositing habit of this tachinid all collecttd material was sent to the Yokohama laboratory for obsershyvation Hamaria incongrua was reared from beetles collected at 68 of these localiti(s on til( 3 islnnds Emelgpnee records from this collected mnterial showed that IIamaria was the moll abundant ill the hilly places cOnled with underbrush in the wnrmer lll(ns Field pamsitizatioll wns excP((lingly YftrinbJe in diffplent spdiol1s unci thl genellll 11Wrnge yely low In 110 locality were morl than 20 percent of till beetks pnrusitized and in only 13 localiti(middots WnS pnmsitizatioll a bovp 5 pCrclll t
Pnlnsi tized b(etles WlI( found ill tlll filhl in ((11 tllll lind westcIll Japan thlougholl t tll( season the highest rates of palUsitization occurring during thf latter part of June and early in July find again the latter part of August At Obuse in western Japan 51 percent oi tlle beetles were parllsitized on Jllly 9 1932 Examination of the beetlfs from the same locnlity coll(cted July 25 nnd 29 rCvealed pnrasitizutions of 23 and 32 pNcpnt respctively which incrensed to 143 percrnt of the beetlls rollcted the last week in August Popillia collected at Takao near Tokyo in enstern Japan on July 3 1932 had n parusitizfl tioD of 11 ppl(ent which dlopprd to 12 flnd 14 percen t in (ollctions made on J tlly 19 nnll 28 lesppctiVCly and rose flgain to 43 p(lcent ill a lot collected August] 9 The differencps in these palnsitization r(gtcorc18 are no doubt tlue to the two generations of llamaJmiddotia I1ppearing in thl field at tlwse times
In alN1S wlHre adults of Popillin japonicn arp prCs(nt in thC fidel over a period of ~ months or longer llam(uia in(ongrua is ab to develop two glneratiom on this host but in arcas where this dOlS not occur flltcrnatc scarnbacid host bectles a Ie lleCPSS1UT for the pnS(lshyvation of this species
6 TECHNICAL BULLETIN 738 U S DEPT OF AGRICCLTGHE
EUlRIXOPSIS J VANA Townsend
In 1923 nnd 1928 a few adults of Eutrixopsis javana were reared at the Riverton laboratory from shipmeuts of beetles parasitized by Oenteiel cinerea forwarded from Sapporo Japan in 1922 and 1927
Although several attempts haYc been made to obtain information on this tachinid in Japan nothing relative to its biology or life history has been found A cardul examination of beetles bearing tllcbinid eggs collected at Snpporo in 1928 revealed that all tlll pupuria conshytained within the dead parasitized beetles wen those of (enteter ~10re than 100000 beetles free from external tacbinid eggs were also examined but not a single Eutrixopsis puparium vas found In 1930 approximately 10000 PopilliCL weI( (oUecteci at 9 otl(r localities on Rokkaido Island ~Jore than half of the beetles bore tachinid eggs and the remainder were free from extemal eggs Luter examination of both pnrnsitizpd and unpuIfisitized beetles revealNl nothing but Centeier puparin
Apparently Eutrilopsis at times parasitizes Popillia in the absence of some other host and may be consici(led of little importance as a fnctor in the control of P j(Lponica in Inpnn
TnOlHOPS CLA lHEN Aldrich
Trophops cla1l8eni was first discoerNl in 1928 whln ~OOO Popillia iaponica brrtlps colllClrd at Tokorozawn neaL Tokyo Japan late in June rcvpaled t parasitization of 12 pCLeent Blrths (ollrcted at this arCH 1 wcek latlL hoe-el did not eontain n single individual
In 1930 while the distribution of and fi(lcl pnrnsitizltioll by (enieter cinerea find lIamaria iIlCOnflla WeIe being inv(stigatNl in XagHno prefedurr westprn Japan Troptops wns again pn(ollutrred in two locnliti(s In 1931 it wus found in sCnrnl ndditiollllllolulitips in the sumc prefpcture The highest pnlHsitiznLion WitS found at Obuse middotwherc collections of Popillia in 19~2 yielded 81 nnd i) pereeut on July 9 Hnd ] 3 rrspecti vlly The field pnrnsitization of hcptllS collected in all loealities however uytrngpd only 14 lwrccnt in that year Of seyrrnl Scarabueidae present in the fi(middotld in these localities Anomala lujocuplea appears to bc the only other beetle a(cpptrd by tlus tachinid as a host Although Popiliia heetles collpded at Toko1ozawa latp in Junc in 1928 we1P found to contnin Trophops laryae no pnrnsitiz((l beetl(s wcre found in NngnTlo prei(ctule brforc July 5 Aftel this date the degne of fipld pnrHsitizntion increnscd until the middle of the month aftrl which it grndllnlly dePlined After the first wepk in August no pnrnsitizcdPopillia could bp found although A rllocuprea coll(cted late in Allgust wpre found to rontain larvue of Trophops Colkctions of berthmiddots made in thr middle of July showed the sex ratio of the attacked brctles to be 61=3 pcrcent mnles nnd 387 perc(nl females
Votes on the L1je llistMy of Trophops claU~flti
Thp tachinid Hy Trophops clauReni is npparCntly l10eturnnl in its habits for it has never he(n obs(ry(d in Ih( filld PlrHsitizcd heetles are uSllnlly found in rather hilly pinelS with dtl1s( undelshygrowth indicating fidel habits similar to those of IIamaxia Hppellteel
7 I~TESTIGATIOXS OF PARASITES OF rAPAXESE BEETLE
examinations of the beetles containing the 1aryue of Trophops showed no traces of external rggs so it is nssuinrd that the femnle deposits living larvae but the exuct mllnnel in which it attacks its host or the manner of penetrution by the 1urva into tll( host heet1( has n(ycr been determined The clrath of the bretle occurs when the purasitc larva has reached the eurly purt of th( thin1 instal TIl( period of huva1 clevrlopmcnt in the host body is similar to that of Hamaria incongrua the ttvemgr time from roHection of the b((tles to pupation of the parasitE being 9 clays nnd(l nOlmnl t(mp(Lll tllles pr(yuiling during midsummer In July nnd August the pupnl stagr in Popiliia COWlS a prriod of 15 to 17 clnys Howrycr in Anomala r1fOC1lprea which werE collected in the latter part of August the pupal peliod was 28 clnys
The rxuct llumb(l of generations of Trophops in (8t(111 Jnpull is not known although there arr at l(ust two a year the first on Popillia und the folloing onr or morf on aailable Popillia nne Anomala lujocuprea In 1982 Tlophops en1llgrd dnring th( rnriy part of August from Popillia b(etl(s that had bern collected in the middle of July und pound1 71tfocllprea colledCd in the latter part of August were also found to contain lnrnlc of this parasite
The stntns of this tnchinid ns a pnmsite of Popillia japonica is son1lwhat similnr to that of [Jamajia incongnw although its known distribution is morE limitrd BrcHusr of its low rnJe of field pamsitishyzntion ns ohs(lwd in its natin habitat in Japan nnd its two or more gruelUtions a yrnr conring it prriod of ndult nctiity of 2 months or long(l the alur of Troplwps clallseni ns a pnrasite of P japonica in the rnitecl Stntes is questionabll
In 1932 1 totnl of 287 pupa ria of TropllOp8 clauseni erl shipPNI to the rnitNI StntrR from which only 16 adults (merged in the spring of 1933
PYHGOTlDAE
ADAPILIA FIA VIfltETA Aldrich
A full account of tIl( lif( history nnd genlral hnbits of Adapsilia jialiwta a parasite of adult Popillia blltlrs in Inclin has heen given in a previous publication (10) In Indin this pyrgotid Il~middot is restricted to the hilly sections that nre eoyered with wieldy Spltc(eI plnntings of pine trees and fnirly dll1sr undergrowth of brush nt an (elltion of about )000 flet It has only ol1r glnlration It year the ndults apppuring in thefilld about thl middll of July Its normal host in India is Popilia cllpricolli8 although P tyanea nnt P macclellandi arl occl1siol1itlly pnrllsitizrcl In parnsitizing th( bp(ll( lhr [(mnle ildapnmiddotZia which is diurnal rrmnins quirs((llt on th( foliag( nfar the feeding br(tlr until thr brrtlr tnkrs flight whr[rupon till fI~ immedishyately pOllnc(s upon thr b(rtlr from nhoH It tlwn quickly insClts its egg through [1 long rrlntiwly slend(r and shnrp ovipositor into the mid-dorsal l(gion of the b((tlr nhdom(n Il(nr the junctllle with thl thorax
Thr work in Indin in H)29 was rrvotc1rl to col1((ting hl(ltlr l1EfLl Sbillong lWei sbi pping 1dapsilia pu pnria obtai nrcl from thpsr hl(tlrs to the Cnited Stt1t(S X0 furthr work was don( with this BI)((ies nftrr 1929 owing to th( dis(ontinuntion of thr work in thttt country
Brrt1Es W(Ire collcet(l(l dllling tIl( period from July 25 to August 18
8 TECHXICAL BULLETIN 738 U S DEPT OF GRICFLTCRE
1929 th( majority being brought in from August 1 to 9 th( p(liod when the synchronization of host and parasite is at its best DlIling this time 553885 beetles we1( brought in by the llum(IOtls l1atic collectors FIOIU these 4310 Adapsilia pupalin were obtained giving a parasitization of 078 percent as compared with 053 in 1928
TIlt manuel of renring uncI shipping wus similnr to thnt (mployed in preyious yenls The beetles were confined in hla cardboard boxes of approximately 1 cubic foot capacity Illld to absorb excess moisture the bottoms w(1e lined with fresh moss Apples were used to f(ed the beetles and were l(phnisbed on alternate dllYS After 12 dnys allliye bC(tles which were (lvid(ntly Ullpn1llsitized W(Ie destroy(d nnd the dead olles placed in opm trnys to allow the Atlapsilia which W(lre still ill the Inrvn1 stngr to pupntt Artel pupntion til( UllCOIlshySUlllNl body rontrnts of th( host wrre rellloved to diminnt( decomshyposition gases that might arise durillg shipmrnt to til( detriment of the pllparia The pupnrin v(1e packed betwNm ltl~nS of fresh sphagnum moss in smull vrlltillLted 1ectnnglllnJ tillS and n number of tlllse en tirely surroUlHlpd by 1 3-illClL layer of fllsh mo (1(
pfldwl in illS(Ctploof woodell box(s for shipment One shipnwnt eonsisting of 3ROO pupnrin was SPIlt to tli( rnited
Stnt(s in 1929 This arrind at its dCstinntion Xonmb(r 9 Illvinp bN1l ell rou tp 42 ditys nt it t(mp(rntue of 40deg to 50 0 F
PARASITES OF TilE LAIVA
nEXIlDAE
DEXIA EXTnTIS Aldrich
Delin 1flltrali tilr most rommon of the ChmltHs( dlxiids Wiltshyfirst found Ilt SUiglll ChosllI in 1922 (9) Owing to its (olllplirntld biology no r(Hring work for the shipnwnt of pnmsitiz(d glU bs to t1H lnit(gtd StatN was aWmpted until 1)21) (10) In that Yenr P()pilli(t jnponic(L grubs whirh do not orcur in Chosen (1( eol1letrd filld shipped from nortlH11l Japan to Fukuoka in Kyusitu soutitNll Jnpitn for pnrnsitizlltion by Duia (mnks fonnumiddottkd from Chos) By thr tinlP n suiJici(nt numb(middot of glflYid f(BmTes Wtl( nvailablr Inte i11 JU11r how(yrr most of the grubs w(re too far d(vdop(d for pnrnshysitiztltion purPOR(S find only 850 pnrnsitizld grubs were shipped to til( enitrd States
In 1926 it WfiS d((idpd to (nlTY on rlaring work in 0hos1 utilizing the nutiY( ChOSlll(RC host grubs This provrd to be very sucressful and OWl 20000 grubs were pamsitizrd flnd shippld to titC rnit((l States in 102(3 lind 1927 B(ctlUSC oJ til( dlLngPl iIlYoiwcl in shipping to the rnited Statts Jive grubs othrl thtlll tilos( of POJlillia iapflnica owing to tIl( risk of 111 rscnpl of thrse sp(cils no l(uring w(lIk WIlf
cHrrild on in Chosen in 1928 but 11 mdhod WIIS dlvCloppd whplpby adult fCmale DIJia could be sitipPllaquo LO Yokollilma SII(clssfully and in 1029 rrnring work wns rnrri(d on til(lC the grll bs of P japof ira being IIs((I tiS hosts
Tabl( 2 shows the slllTinl of ndult f(malCs of [Jain middot(lltr(1Ii forshywllrd((l from SlIig(Il Chospn to YokohfilJ11L in 1029 Hnl 1930 til( yciUS that rCaring work WilS el1JTi(d OIl TIle I1dults PI( ronfillNI in thl shipping containl1s which WCI( similm to thos llIlployltd ill transporting Scoliidne npprox11llat(ly 7 days from the timr o( (oll(eshytion to the tim( of removaL
IXVESTIGATIOXS OF PARASITES OF TAPANK~ BEETLE 9
TABLE 2-SuTviLal of adult females of DeJia ventralis shipped from 81tigcn Chosen 10 Yokohama Japa11in 1929 and 1930
Adults Adults surllingYear Generntion shipped shipment
Numb lrumber PercentI
1929_ bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull _ FirsLbullbullbullbullbullbullbullbullbullbullbullbull _bull __ bull __ bullbull_1 063 552 833 1921 __ bullbullbullbullbull ______ bull __ bullbull ____ lhird____ __ 2281 1800 831 1930 __ _bullbullbullbullbullbull __ bull __ First _______ __ ~9() 6S- 68 1910_ _ __ _____ bull_____ __ 1 erond_____ __ __ 2051 14il 720 1930bullbullbull __ bullbullbullbull __ Third __ 2483 2008 809
Total __ __ __ ___ 8368 t 661 ]----()-1 I
It was extremrly difficult to obtain Popillia grubs in any numbers in the --icinity of Yokohamn for rearing work with Deria but this was OYerC0111( by collecting ancl shipping grubs as IHeclNI from Hokshykaido in nortlltrn Japall to Yokohallla n procedure which proyrd to be yery satisfactory and inrxplnsiYe ~0 rearing work was attempted with the s(cond gel1lrntion of DeJia in 1929 oing to the impossibility of s(curing u sufficient number of grubs in IIokkuido during August This was the YNlr of grub scarcity in northern Japall vhen less than 1 percent of the oYCrwintering grubs huying a 2-y(middotnr lifc cyclc (le still in the brynl stage at thn t time of the yeal
The ptlrasitizNl grubs obtain(d tluough parasitization with the first and s(concl g(l1Plntions of DU(L in June nnd August lPspCctiYely wele shipped to thpCnitltC1 Stat(s immpdiatrly aftC1 btinf pnra~itizcd DcYClopment of the parasites took pla(e P11 route nnd the adults emrTgecl sho1tl)- uiter nrriyal at tlHlT destination Grubs containing tbe hibernnting ln1yue of De ia obtuined through pnrasitizatiOIl with the third-g(lHlntion 1Ii(s in 8(ptp111her nnd Octol)(r Pre kPpt OY(1 wintpr at 10kohi1111tl in n wlll-crain(c plot (oY(INI with sod The winttr lllortnlity of these grubs was rnthel high ns only 171 and HL9 pcrcent werc nIl( when dug up [or shipnlPnt in the springs of 1930 lllHl 1931 rlspertiwly
Rwring and Shipping Jfpoundt7wds
The rcuring work with lJJ(l l(fltl(llj1 was accomplisll(d hy using a bnmiddotpding-jnl 111Cthod simillll to that employrd in ]Jle-iolls y(nrs Approximately 50 grubs (Ie placfll in the lonrhalf of a l-qualmiddott glnss jar with soil A fin(-IJ1Psh SCIepn cngp contllining 5 or () lJ(tia f(malp WIlS fittNI into this jur lind rPRtpd on til( soil nnd through this scnPI1 into the soil til( dlpositcd lalTHP pnsNI in palch of the host gruhs 1llP grubs wpl(~ (xtunilHd pneh chty Hlld thORP showing thf pnrnsit( nmiddotpjlntol~ funlwl (1( ((monel nne Tplacpd with frlsh g-rub The flips which Pl( fpd IPg III ll[l with it solution of sunr und watN linmiddotd f[om 3 middot(middotCk tc) J IJ1fll1th in til( cage nnd pnrnitizld flom a5 to )0 pP[cpnt of tlH grub Iach dnv
In pucking the pnrnsitizplt1 gruh 101 shipnwnt iin-lillpd hoxls 1 foot squnrp und Ii inch(s dNp W(IP wwd Each l)()x contnilpd 5 cro-5lction s(ts RPparutd b lin pintp ploviding it COll1pnltllHnt of nhout 1 (lbic inch foJ Pilch of jOIl glubs Fol shipllllllt 2 of thpse hoxps (1( packlrl togtlwl in n (Tong woodpl1 (nsp TIl( soil in which thp gru bs (n pu(k(d had hren thorough Iy fumign tld wi t h (arbon (hsulfid( to (limirmtC any otlw1 inlcl lllntrrial which might htwe
23-13-1----40-2
]0 TECH~ICAL BULLETIN 738 U S DEPT OF AGRICULTURE
been present In table 3 is a summary of shipments of palasitized grubs and the percentages of parasite emcrgenee after arrivul in thl United Stntls
TABLE 3-SlImmary oj shipments oj Popillia japonica grubs l)ara~itizcd willi Deda Ientralis Jrom Japan to the United States 1929-31
Year Contents oC shipment Grubs Parnsites emergingship(led-------1--------------- ---- ______ lVumber lVlLmber I Pentnt
19211-bullbullbullbullbullbullbullbullbullbull_bullbull Grubs pnmsiti1ed with flrstgenerntion Dezia ~ ~500 ~51 1000 1930_bullbullbull __ bull __ bullbullbullbullbull __ Overwintered grubs parnsitized with thirdmiddot 3--
gcnemtionDaiu ~~_~ __ ~~ ___ _ ___ ~ 3000 II 126 1930bullbullbull ___bullbullbullbullbull __bull ___ Grubs pamsitized with lirst-enerntion Detia~~ 8000 1504 18S 1930_____________ ~ _ __ Orubs pllrasitized with ~econd-generntion Drtia 7000 l42J I 203 1930____ __ ~ ~ ~ _ GrUbs pllmsiti1ed with second-genemtion 2500 399 16 (f
Dufu II1931___________ Oerwintered grubs paraSitized with third- 4000 345 8 ( generation Duia I
I 1--------- shyTotaJorneruge~_i- ___ bullbullbullbull __ _ _______ bull ___ ~ ~ ~)(JOOO I 4499middot 155 f i i
Owing to the establishment of Diria lentrali~ in tIl( United Stutes from which materinl for lccolonizlltion was availabll no shipments of this pfllflsitl Wlre made aftlr 1931
PnOSENA SIBII1TA (Fabricins)
Thl dexiid fly Prosena sibi7ita middotliich IS glllclfllly distrihut(d throughout till Asiatic and ~IalaYlln Hegions is an importnnt pilll1shy
sill of PopWia iaponica in Kolth(1l1 Japan It hns olle glllelation a YfIU ami is tllpubl( of palllsitizing and dfwloping Oll numplou sppcils of sealt1bnpid grubs of suitnblr size The life history nnd ililbits of this dexiicl huve been fullv dPidt with in previolls publieashytions (9 10) bull
The shipments from 1923 to H)27 inclusi-e cOl1[-iistN[ of fitldshycolkrtfd grubs of Popillia iap(mit(l whieh showld a getHrnI fitld parflsitization of 10 to 15 percpnt Becaust of till impossibility of detCrting the plesenee of the pUlllsite latnt whieh is nttuchNI to tlH main trnelwal trunk of the host all collpcted glUbs W(I( shipp((L Tllf grubs (re dug in Hoklmido rudy in the spring lind imnHdilltely forwarded to the rnited States arriving thete shortly beforp tll( emlIgenee of the parasites Although this method proed sntisshyfactory the trllnsportntioll costs wPle (xtrCll1(ly high nnd in 1927 it was dfcided to palllsitize till grubs nrtifjeiall)- by 11 mltthod that hnd bCll1 laquo(eloped in 1921 This ms aecomplished by dissecting til( first-stag( larval from tIle oVlsacs of the glUyid femuks and trilllSshy
ferring them to thC host grubs by mcans of a fine blllsh To instlll penetration by the parnsites the grubs with til( smal lnlYllc wIe vlaeed in cclIs of suitable size in wooden bloek fotmiddot it petiot of 24 hours brron being plaecd in thc field whftP tllr oYClwintrlld
Late in the summer of 1928 40596 grubs W(Il artificially PUIllshysitized in 1l0rthCrTJ Japan llnd placed in a prcpnred soil plot w(11 drained coered with sod and surrounded to a depth of 18 inch(s by a fine-mesh screen to pr(vent the gnlbs from wumlcring nwny T11(s( grubs were dug up carly in May 1929 and 19440 were nliv( of whieh 19000 weIpound shipped to the U ni ted Stat(s
In 1929 it was impossibl( to seure sufficient grubs for inoeulatioll work Consequently it was necessary to colleet Prosena femulCs and
IXETrGATIOXS OF PARASITES 01 rAlANFJSFJ DEETLB 11
ship them to the Yokohnma laboTfitory und carryon palasitization work there with available current-year grubs from that vicinity Only 12 p(rcent of the females were alive on arrival at Yokohama Tlus high mortality WitS due to the extremely hot weather prevailing at the time of shipment Out of 364 females received alive 159 contained Inrvae sufficiently developed for this work During the latter part of August 6300 grubs were parasitized and placed in hibernation qunrters As e~perienced with the overwinter~d grubs parasitized with Dexia 1entralis the wintPI mortality was (xceedingly high in this material and only 900 were alive wh(n dug up the followshying spring These with the 19000 shipped in 1929 together rith the 29000 of DeJia tentralis make a total of 48900 grubs containing dexiids thnt were shipped to the United States during the period covcl(d by this bulletin The method of pl1cking tIl( pnrasitized grubs for shipment was the sam( I1S that described for shipment of grubs parnsitized with D ~entrais
No shipments of Prosena sibirita wer( mude aftN 1930 ns this parasite was then established in the Vnit((1 States
SCOLIIDAE
During the course of the investigations from 1920 to 1933 inclusive on the pnrasites of Popillia and related scarabaeid b(etl(s in the Orient a huge number of Scoliidne principally of the genus Tiphia were found and intensively studied ~fost of thtsP species which were consid(recl to be ntw have since been des(ribtd (2 18 19 2021)
In table 4 is presented the 1llown distribution in Tnpllll ChOS(D China Indin nncI Taiwan of all species of Tiphia real(d or shipped sinet the initilltion of the work Of the 29 sp(ei(s list(ci in this tablr 8 nre known to occur ill Japan Chosrn and China 2 in Jupnll nnd ChOS(Il and 1 in ChOS(1l and Chinn No doubt furtlH1 innstigtltions will show a much wider dissemination throughout the Orient Of the species mentioned in this bulletin only liphia bisfculata nnd T tegitiplaga purnsites of Anomala allld occurring in southern Jnpnn have 2 distinct generations a year
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
j
OQ 13
0 0 lts ~
lt ~ a ~ ~ ~ ~
gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
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4 TECHNICAL ll1LLETIX 738 L S DEPT OF AGlUCrLTCHE
TACHINIDAE
CENlETER CIXEREA Aldrich
The biology and habits of Ceniefl7 cinerea the m08t cfTectin palllsite of adult beetles of Popillia iaponica in northcrn Japnn I111s bccll fully (leaH with in previous publications (D 10) Thc work during the period 1929-33 consisted in the inYCstigntion of llCV nlen8 in Tnpnn to determine thc relatiye abundancc and dpgrp( of fiplcl pnmsitizntion and in the collection of pnrnsitizpd i)(etles for the shipll1lnt of puparin of C cinerea to the United States
IUYfstigations on the islnnd of Hokknido in northpll1 Tnp~lIl reYCaled that Centeter cinerea was well distributed in pmcticaIly all the localities vhere Popillia beetles were present The nTcmge parasitization in all the localities innstigated was 372 percent during the hltter part of July and early part of August in 1931 No doubt this anrngr would have been considerably higher hnd the obscrvutions lWln made cudirr in the senson since preyious records at Sapporo on tIll Silme island have shown that the parasitization drops of1 very sl1ddenlmiddot Ilfter the first part of August On the island of Honshu just south of Hokkaido the presencp of C cinerea in different sections was distinctI -ariable nne the degree of parasitization considerably lower as Contrnstpd with that found in Hokkaido In no locality wpre more than j) pprcent of the beetles pnrasitizecl The highest Utes of pnrnsitizn tioll wrre found in the hilly and mountainous sections of the northprn and western portion of the island where the win telS arp sCYerl Hnd SHOW is present on the ground throughout the wintCr In thC lowlnnds of the pastern portion of the island where the winters arp mild und little SIlOW
)ccurs beptlc parasitization was found to be considCrubl- lowpr and in the section south of Yokohama no trace of C cinerw WI1S found
1Yhile the distribution of Hamaxia incongwa was blillg innstigatNI in Kyushu in southern Japan in 1032 beCtlps baring eggs sil11ilur to those of Ce7~teter were colIeetNI near Ino and Kobayashi in the mounshytains at the northern end of the island mergrnc( of til( pnmsitls the following year proYed them to be C cinerea This makes thr third island in Japan on whieh 0 cinerea is known to occur and (xUnels the known southfrn range much beyond that previously r((ordpd
Popillia bpctlts beaTing Centeter eggs wCre collrctCd for llI)ring and shipment in ] 929 1930 und 1933 Approximattly 85 p(reent of tllC parasitized beetles produced Centeter- pupa ria During thrsr ylturs 33663 puparia were sbipped to the UnitCd StatCs The 1933 shipshymtnts consisting of 6163 puparia were from brrtlCs (0llpC t ((I in the yicinity of rtsonomiya and Tachikawil T1Enr Tokyo The (entfler ndults occurring in these lo(alities are prpsent in th( fipld tllp lattpr Pfilt of Junp and early in JlIl- or approximntrly 1 month Nlrlipr thun those at Sapporo in northCm Jnpan from whieh all prpyiolls hipnlPnts of pupnrin l11atprial have originat((1 fn virw of this and thl (OITPshysponding COITllatioll with th( timc of rl11prgPlw( of thp hot ( cinerea from this section mny prOY( to be hpttEl sYllchronizpd ill timp ()[ mCrgpnee with POJ1illia japonica in the Cnitcd Statls
The mCtboltis of (ollpcting nnt lpaling WltJ( the samp a thosC rl11shyployltltl in pl(iolls ypars Tn plPparing tilpound pllpnritl [or hipnlCnt great (nre nlS exCrciseu in eliminating as much of the bpptip hody as possiblp
5 IXYESTlGATIOXS OF PARASITES OF JAPAXESE BEETLE
HAMAXIA INCONGRtA Vnlklr
llam(llia inconglua a pmusite of the adults of many species of Scarabutidae is widely distributed throughout the Asiatic and IVlalaYfill Regions find in various tropicn countries In Japan where it occurs as a parasite of Popillia japonica it has two and possibly thrre genErntions a year Bping of crepuscular habitJ it is active mostly during the carlr morning hours and in the cvening Although the lxuct mlthod of parasitization is not known it is prlSUDlN that thc femult dlposits fully deClopN lalYap lither on the host beltle or on the llrnrby foliagr and that tlipse enter the beetle through thc softer portions of t111 body
Although no shipments of IJama11aincongrua have bern made to the Lnitrd StlltlS since 1928 obsllyations on distribution and degree of field pHrnsitizatioIl werc carrid on in Japan during th period 1929-33
During 1930 1931 and H)3~ onl 200000 Popillia beetlls (1(
collected nt 108 localitis lepnsenting a wide range of climatic tlnll enviroIlmeJltal conditions 011 the islnnds of Honshu Sbikoku and Kyushu in lapan These were collected from the middle of Junc to the end of August nod in some instances eol1ections were made at weekly internlls Since it is impossible to distinguish the pnrasitizpltl beetles in the field owing to the lnrvipositing habit of this tachinid all collecttd material was sent to the Yokohama laboratory for obsershyvation Hamaria incongrua was reared from beetles collected at 68 of these localiti(s on til( 3 islnnds Emelgpnee records from this collected mnterial showed that IIamaria was the moll abundant ill the hilly places cOnled with underbrush in the wnrmer lll(ns Field pamsitizatioll wns excP((lingly YftrinbJe in diffplent spdiol1s unci thl genellll 11Wrnge yely low In 110 locality were morl than 20 percent of till beetks pnrusitized and in only 13 localiti(middots WnS pnmsitizatioll a bovp 5 pCrclll t
Pnlnsi tized b(etles WlI( found ill tlll filhl in ((11 tllll lind westcIll Japan thlougholl t tll( season the highest rates of palUsitization occurring during thf latter part of June and early in July find again the latter part of August At Obuse in western Japan 51 percent oi tlle beetles were parllsitized on Jllly 9 1932 Examination of the beetlfs from the same locnlity coll(cted July 25 nnd 29 rCvealed pnrasitizutions of 23 and 32 pNcpnt respctively which incrensed to 143 percrnt of the beetlls rollcted the last week in August Popillia collected at Takao near Tokyo in enstern Japan on July 3 1932 had n parusitizfl tioD of 11 ppl(ent which dlopprd to 12 flnd 14 percen t in (ollctions made on J tlly 19 nnll 28 lesppctiVCly and rose flgain to 43 p(lcent ill a lot collected August] 9 The differencps in these palnsitization r(gtcorc18 are no doubt tlue to the two generations of llamaJmiddotia I1ppearing in thl field at tlwse times
In alN1S wlHre adults of Popillin japonicn arp prCs(nt in thC fidel over a period of ~ months or longer llam(uia in(ongrua is ab to develop two glneratiom on this host but in arcas where this dOlS not occur flltcrnatc scarnbacid host bectles a Ie lleCPSS1UT for the pnS(lshyvation of this species
6 TECHNICAL BULLETIN 738 U S DEPT OF AGRICCLTGHE
EUlRIXOPSIS J VANA Townsend
In 1923 nnd 1928 a few adults of Eutrixopsis javana were reared at the Riverton laboratory from shipmeuts of beetles parasitized by Oenteiel cinerea forwarded from Sapporo Japan in 1922 and 1927
Although several attempts haYc been made to obtain information on this tachinid in Japan nothing relative to its biology or life history has been found A cardul examination of beetles bearing tllcbinid eggs collected at Snpporo in 1928 revealed that all tlll pupuria conshytained within the dead parasitized beetles wen those of (enteter ~10re than 100000 beetles free from external tacbinid eggs were also examined but not a single Eutrixopsis puparium vas found In 1930 approximately 10000 PopilliCL weI( (oUecteci at 9 otl(r localities on Rokkaido Island ~Jore than half of the beetles bore tachinid eggs and the remainder were free from extemal eggs Luter examination of both pnrnsitizpd and unpuIfisitized beetles revealNl nothing but Centeier puparin
Apparently Eutrilopsis at times parasitizes Popillia in the absence of some other host and may be consici(led of little importance as a fnctor in the control of P j(Lponica in Inpnn
TnOlHOPS CLA lHEN Aldrich
Trophops cla1l8eni was first discoerNl in 1928 whln ~OOO Popillia iaponica brrtlps colllClrd at Tokorozawn neaL Tokyo Japan late in June rcvpaled t parasitization of 12 pCLeent Blrths (ollrcted at this arCH 1 wcek latlL hoe-el did not eontain n single individual
In 1930 while the distribution of and fi(lcl pnrnsitizltioll by (enieter cinerea find lIamaria iIlCOnflla WeIe being inv(stigatNl in XagHno prefedurr westprn Japan Troptops wns again pn(ollutrred in two locnliti(s In 1931 it wus found in sCnrnl ndditiollllllolulitips in the sumc prefpcture The highest pnlHsitiznLion WitS found at Obuse middotwherc collections of Popillia in 19~2 yielded 81 nnd i) pereeut on July 9 Hnd ] 3 rrspecti vlly The field pnrnsitization of hcptllS collected in all loealities however uytrngpd only 14 lwrccnt in that year Of seyrrnl Scarabueidae present in the fi(middotld in these localities Anomala lujocuplea appears to bc the only other beetle a(cpptrd by tlus tachinid as a host Although Popiliia heetles collpded at Toko1ozawa latp in Junc in 1928 we1P found to contnin Trophops laryae no pnrnsitiz((l beetl(s wcre found in NngnTlo prei(ctule brforc July 5 Aftel this date the degne of fipld pnrHsitizntion increnscd until the middle of the month aftrl which it grndllnlly dePlined After the first wepk in August no pnrnsitizcdPopillia could bp found although A rllocuprea coll(cted late in Allgust wpre found to rontain larvue of Trophops Colkctions of berthmiddots made in thr middle of July showed the sex ratio of the attacked brctles to be 61=3 pcrcent mnles nnd 387 perc(nl females
Votes on the L1je llistMy of Trophops claU~flti
Thp tachinid Hy Trophops clauReni is npparCntly l10eturnnl in its habits for it has never he(n obs(ry(d in Ih( filld PlrHsitizcd heetles are uSllnlly found in rather hilly pinelS with dtl1s( undelshygrowth indicating fidel habits similar to those of IIamaxia Hppellteel
7 I~TESTIGATIOXS OF PARASITES OF rAPAXESE BEETLE
examinations of the beetles containing the 1aryue of Trophops showed no traces of external rggs so it is nssuinrd that the femnle deposits living larvae but the exuct mllnnel in which it attacks its host or the manner of penetrution by the 1urva into tll( host heet1( has n(ycr been determined The clrath of the bretle occurs when the purasitc larva has reached the eurly purt of th( thin1 instal TIl( period of huva1 clevrlopmcnt in the host body is similar to that of Hamaria incongrua the ttvemgr time from roHection of the b((tles to pupation of the parasitE being 9 clays nnd(l nOlmnl t(mp(Lll tllles pr(yuiling during midsummer In July nnd August the pupnl stagr in Popiliia COWlS a prriod of 15 to 17 clnys Howrycr in Anomala r1fOC1lprea which werE collected in the latter part of August the pupal peliod was 28 clnys
The rxuct llumb(l of generations of Trophops in (8t(111 Jnpull is not known although there arr at l(ust two a year the first on Popillia und the folloing onr or morf on aailable Popillia nne Anomala lujocuprea In 1982 Tlophops en1llgrd dnring th( rnriy part of August from Popillia b(etl(s that had bern collected in the middle of July und pound1 71tfocllprea colledCd in the latter part of August were also found to contain lnrnlc of this parasite
The stntns of this tnchinid ns a pnmsite of Popillia japonica is son1lwhat similnr to that of [Jamajia incongnw although its known distribution is morE limitrd BrcHusr of its low rnJe of field pamsitishyzntion ns ohs(lwd in its natin habitat in Japan nnd its two or more gruelUtions a yrnr conring it prriod of ndult nctiity of 2 months or long(l the alur of Troplwps clallseni ns a pnrasite of P japonica in the rnitecl Stntes is questionabll
In 1932 1 totnl of 287 pupa ria of TropllOp8 clauseni erl shipPNI to the rnitNI StntrR from which only 16 adults (merged in the spring of 1933
PYHGOTlDAE
ADAPILIA FIA VIfltETA Aldrich
A full account of tIl( lif( history nnd genlral hnbits of Adapsilia jialiwta a parasite of adult Popillia blltlrs in Inclin has heen given in a previous publication (10) In Indin this pyrgotid Il~middot is restricted to the hilly sections that nre eoyered with wieldy Spltc(eI plnntings of pine trees and fnirly dll1sr undergrowth of brush nt an (elltion of about )000 flet It has only ol1r glnlration It year the ndults apppuring in thefilld about thl middll of July Its normal host in India is Popilia cllpricolli8 although P tyanea nnt P macclellandi arl occl1siol1itlly pnrllsitizrcl In parnsitizing th( bp(ll( lhr [(mnle ildapnmiddotZia which is diurnal rrmnins quirs((llt on th( foliag( nfar the feeding br(tlr until thr brrtlr tnkrs flight whr[rupon till fI~ immedishyately pOllnc(s upon thr b(rtlr from nhoH It tlwn quickly insClts its egg through [1 long rrlntiwly slend(r and shnrp ovipositor into the mid-dorsal l(gion of the b((tlr nhdom(n Il(nr the junctllle with thl thorax
Thr work in Indin in H)29 was rrvotc1rl to col1((ting hl(ltlr l1EfLl Sbillong lWei sbi pping 1dapsilia pu pnria obtai nrcl from thpsr hl(tlrs to the Cnited Stt1t(S X0 furthr work was don( with this BI)((ies nftrr 1929 owing to th( dis(ontinuntion of thr work in thttt country
Brrt1Es W(Ire collcet(l(l dllling tIl( period from July 25 to August 18
8 TECHXICAL BULLETIN 738 U S DEPT OF GRICFLTCRE
1929 th( majority being brought in from August 1 to 9 th( p(liod when the synchronization of host and parasite is at its best DlIling this time 553885 beetles we1( brought in by the llum(IOtls l1atic collectors FIOIU these 4310 Adapsilia pupalin were obtained giving a parasitization of 078 percent as compared with 053 in 1928
TIlt manuel of renring uncI shipping wus similnr to thnt (mployed in preyious yenls The beetles were confined in hla cardboard boxes of approximately 1 cubic foot capacity Illld to absorb excess moisture the bottoms w(1e lined with fresh moss Apples were used to f(ed the beetles and were l(phnisbed on alternate dllYS After 12 dnys allliye bC(tles which were (lvid(ntly Ullpn1llsitized W(Ie destroy(d nnd the dead olles placed in opm trnys to allow the Atlapsilia which W(lre still ill the Inrvn1 stngr to pupntt Artel pupntion til( UllCOIlshySUlllNl body rontrnts of th( host wrre rellloved to diminnt( decomshyposition gases that might arise durillg shipmrnt to til( detriment of the pllparia The pupnrin v(1e packed betwNm ltl~nS of fresh sphagnum moss in smull vrlltillLted 1ectnnglllnJ tillS and n number of tlllse en tirely surroUlHlpd by 1 3-illClL layer of fllsh mo (1(
pfldwl in illS(Ctploof woodell box(s for shipment One shipnwnt eonsisting of 3ROO pupnrin was SPIlt to tli( rnited
Stnt(s in 1929 This arrind at its dCstinntion Xonmb(r 9 Illvinp bN1l ell rou tp 42 ditys nt it t(mp(rntue of 40deg to 50 0 F
PARASITES OF TilE LAIVA
nEXIlDAE
DEXIA EXTnTIS Aldrich
Delin 1flltrali tilr most rommon of the ChmltHs( dlxiids Wiltshyfirst found Ilt SUiglll ChosllI in 1922 (9) Owing to its (olllplirntld biology no r(Hring work for the shipnwnt of pnmsitiz(d glU bs to t1H lnit(gtd StatN was aWmpted until 1)21) (10) In that Yenr P()pilli(t jnponic(L grubs whirh do not orcur in Chosen (1( eol1letrd filld shipped from nortlH11l Japan to Fukuoka in Kyusitu soutitNll Jnpitn for pnrnsitizlltion by Duia (mnks fonnumiddottkd from Chos) By thr tinlP n suiJici(nt numb(middot of glflYid f(BmTes Wtl( nvailablr Inte i11 JU11r how(yrr most of the grubs w(re too far d(vdop(d for pnrnshysitiztltion purPOR(S find only 850 pnrnsitizld grubs were shipped to til( enitrd States
In 1926 it WfiS d((idpd to (nlTY on rlaring work in 0hos1 utilizing the nutiY( ChOSlll(RC host grubs This provrd to be very sucressful and OWl 20000 grubs were pamsitizrd flnd shippld to titC rnit((l States in 102(3 lind 1927 B(ctlUSC oJ til( dlLngPl iIlYoiwcl in shipping to the rnited Statts Jive grubs othrl thtlll tilos( of POJlillia iapflnica owing to tIl( risk of 111 rscnpl of thrse sp(cils no l(uring w(lIk WIlf
cHrrild on in Chosen in 1928 but 11 mdhod WIIS dlvCloppd whplpby adult fCmale DIJia could be sitipPllaquo LO Yokollilma SII(clssfully and in 1029 rrnring work wns rnrri(d on til(lC the grll bs of P japof ira being IIs((I tiS hosts
Tabl( 2 shows the slllTinl of ndult f(malCs of [Jain middot(lltr(1Ii forshywllrd((l from SlIig(Il Chospn to YokohfilJ11L in 1029 Hnl 1930 til( yciUS that rCaring work WilS el1JTi(d OIl TIle I1dults PI( ronfillNI in thl shipping containl1s which WCI( similm to thos llIlployltd ill transporting Scoliidne npprox11llat(ly 7 days from the timr o( (oll(eshytion to the tim( of removaL
IXVESTIGATIOXS OF PARASITES OF TAPANK~ BEETLE 9
TABLE 2-SuTviLal of adult females of DeJia ventralis shipped from 81tigcn Chosen 10 Yokohama Japa11in 1929 and 1930
Adults Adults surllingYear Generntion shipped shipment
Numb lrumber PercentI
1929_ bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull _ FirsLbullbullbullbullbullbullbullbullbullbullbullbull _bull __ bull __ bullbull_1 063 552 833 1921 __ bullbullbullbullbull ______ bull __ bullbull ____ lhird____ __ 2281 1800 831 1930 __ _bullbullbullbullbullbull __ bull __ First _______ __ ~9() 6S- 68 1910_ _ __ _____ bull_____ __ 1 erond_____ __ __ 2051 14il 720 1930bullbullbull __ bullbullbullbull __ Third __ 2483 2008 809
Total __ __ __ ___ 8368 t 661 ]----()-1 I
It was extremrly difficult to obtain Popillia grubs in any numbers in the --icinity of Yokohamn for rearing work with Deria but this was OYerC0111( by collecting ancl shipping grubs as IHeclNI from Hokshykaido in nortlltrn Japall to Yokohallla n procedure which proyrd to be yery satisfactory and inrxplnsiYe ~0 rearing work was attempted with the s(cond gel1lrntion of DeJia in 1929 oing to the impossibility of s(curing u sufficient number of grubs in IIokkuido during August This was the YNlr of grub scarcity in northern Japall vhen less than 1 percent of the oYCrwintering grubs huying a 2-y(middotnr lifc cyclc (le still in the brynl stage at thn t time of the yeal
The ptlrasitizNl grubs obtain(d tluough parasitization with the first and s(concl g(l1Plntions of DU(L in June nnd August lPspCctiYely wele shipped to thpCnitltC1 Stat(s immpdiatrly aftC1 btinf pnra~itizcd DcYClopment of the parasites took pla(e P11 route nnd the adults emrTgecl sho1tl)- uiter nrriyal at tlHlT destination Grubs containing tbe hibernnting ln1yue of De ia obtuined through pnrasitizatiOIl with the third-g(lHlntion 1Ii(s in 8(ptp111her nnd Octol)(r Pre kPpt OY(1 wintpr at 10kohi1111tl in n wlll-crain(c plot (oY(INI with sod The winttr lllortnlity of these grubs was rnthel high ns only 171 and HL9 pcrcent werc nIl( when dug up [or shipnlPnt in the springs of 1930 lllHl 1931 rlspertiwly
Rwring and Shipping Jfpoundt7wds
The rcuring work with lJJ(l l(fltl(llj1 was accomplisll(d hy using a bnmiddotpding-jnl 111Cthod simillll to that employrd in ]Jle-iolls y(nrs Approximately 50 grubs (Ie placfll in the lonrhalf of a l-qualmiddott glnss jar with soil A fin(-IJ1Psh SCIepn cngp contllining 5 or () lJ(tia f(malp WIlS fittNI into this jur lind rPRtpd on til( soil nnd through this scnPI1 into the soil til( dlpositcd lalTHP pnsNI in palch of the host gruhs 1llP grubs wpl(~ (xtunilHd pneh chty Hlld thORP showing thf pnrnsit( nmiddotpjlntol~ funlwl (1( ((monel nne Tplacpd with frlsh g-rub The flips which Pl( fpd IPg III ll[l with it solution of sunr und watN linmiddotd f[om 3 middot(middotCk tc) J IJ1fll1th in til( cage nnd pnrnitizld flom a5 to )0 pP[cpnt of tlH grub Iach dnv
In pucking the pnrnsitizplt1 gruh 101 shipnwnt iin-lillpd hoxls 1 foot squnrp und Ii inch(s dNp W(IP wwd Each l)()x contnilpd 5 cro-5lction s(ts RPparutd b lin pintp ploviding it COll1pnltllHnt of nhout 1 (lbic inch foJ Pilch of jOIl glubs Fol shipllllllt 2 of thpse hoxps (1( packlrl togtlwl in n (Tong woodpl1 (nsp TIl( soil in which thp gru bs (n pu(k(d had hren thorough Iy fumign tld wi t h (arbon (hsulfid( to (limirmtC any otlw1 inlcl lllntrrial which might htwe
23-13-1----40-2
]0 TECH~ICAL BULLETIN 738 U S DEPT OF AGRICULTURE
been present In table 3 is a summary of shipments of palasitized grubs and the percentages of parasite emcrgenee after arrivul in thl United Stntls
TABLE 3-SlImmary oj shipments oj Popillia japonica grubs l)ara~itizcd willi Deda Ientralis Jrom Japan to the United States 1929-31
Year Contents oC shipment Grubs Parnsites emergingship(led-------1--------------- ---- ______ lVumber lVlLmber I Pentnt
19211-bullbullbullbullbullbullbullbullbullbull_bullbull Grubs pnmsiti1ed with flrstgenerntion Dezia ~ ~500 ~51 1000 1930_bullbullbull __ bull __ bullbullbullbullbull __ Overwintered grubs parnsitized with thirdmiddot 3--
gcnemtionDaiu ~~_~ __ ~~ ___ _ ___ ~ 3000 II 126 1930bullbullbull ___bullbullbullbullbull __bull ___ Grubs pamsitized with lirst-enerntion Detia~~ 8000 1504 18S 1930_____________ ~ _ __ Orubs pllrasitized with ~econd-generntion Drtia 7000 l42J I 203 1930____ __ ~ ~ ~ _ GrUbs pllmsiti1ed with second-genemtion 2500 399 16 (f
Dufu II1931___________ Oerwintered grubs paraSitized with third- 4000 345 8 ( generation Duia I
I 1--------- shyTotaJorneruge~_i- ___ bullbullbullbull __ _ _______ bull ___ ~ ~ ~)(JOOO I 4499middot 155 f i i
Owing to the establishment of Diria lentrali~ in tIl( United Stutes from which materinl for lccolonizlltion was availabll no shipments of this pfllflsitl Wlre made aftlr 1931
PnOSENA SIBII1TA (Fabricins)
Thl dexiid fly Prosena sibi7ita middotliich IS glllclfllly distrihut(d throughout till Asiatic and ~IalaYlln Hegions is an importnnt pilll1shy
sill of PopWia iaponica in Kolth(1l1 Japan It hns olle glllelation a YfIU ami is tllpubl( of palllsitizing and dfwloping Oll numplou sppcils of sealt1bnpid grubs of suitnblr size The life history nnd ililbits of this dexiicl huve been fullv dPidt with in previolls publieashytions (9 10) bull
The shipments from 1923 to H)27 inclusi-e cOl1[-iistN[ of fitldshycolkrtfd grubs of Popillia iap(mit(l whieh showld a getHrnI fitld parflsitization of 10 to 15 percpnt Becaust of till impossibility of detCrting the plesenee of the pUlllsite latnt whieh is nttuchNI to tlH main trnelwal trunk of the host all collpcted glUbs W(I( shipp((L Tllf grubs (re dug in Hoklmido rudy in the spring lind imnHdilltely forwarded to the rnited States arriving thete shortly beforp tll( emlIgenee of the parasites Although this method proed sntisshyfactory the trllnsportntioll costs wPle (xtrCll1(ly high nnd in 1927 it was dfcided to palllsitize till grubs nrtifjeiall)- by 11 mltthod that hnd bCll1 laquo(eloped in 1921 This ms aecomplished by dissecting til( first-stag( larval from tIle oVlsacs of the glUyid femuks and trilllSshy
ferring them to thC host grubs by mcans of a fine blllsh To instlll penetration by the parnsites the grubs with til( smal lnlYllc wIe vlaeed in cclIs of suitable size in wooden bloek fotmiddot it petiot of 24 hours brron being plaecd in thc field whftP tllr oYClwintrlld
Late in the summer of 1928 40596 grubs W(Il artificially PUIllshysitized in 1l0rthCrTJ Japan llnd placed in a prcpnred soil plot w(11 drained coered with sod and surrounded to a depth of 18 inch(s by a fine-mesh screen to pr(vent the gnlbs from wumlcring nwny T11(s( grubs were dug up carly in May 1929 and 19440 were nliv( of whieh 19000 weIpound shipped to the U ni ted Stat(s
In 1929 it was impossibl( to seure sufficient grubs for inoeulatioll work Consequently it was necessary to colleet Prosena femulCs and
IXETrGATIOXS OF PARASITES 01 rAlANFJSFJ DEETLB 11
ship them to the Yokohnma laboTfitory und carryon palasitization work there with available current-year grubs from that vicinity Only 12 p(rcent of the females were alive on arrival at Yokohama Tlus high mortality WitS due to the extremely hot weather prevailing at the time of shipment Out of 364 females received alive 159 contained Inrvae sufficiently developed for this work During the latter part of August 6300 grubs were parasitized and placed in hibernation qunrters As e~perienced with the overwinter~d grubs parasitized with Dexia 1entralis the wintPI mortality was (xceedingly high in this material and only 900 were alive wh(n dug up the followshying spring These with the 19000 shipped in 1929 together rith the 29000 of DeJia tentralis make a total of 48900 grubs containing dexiids thnt were shipped to the United States during the period covcl(d by this bulletin The method of pl1cking tIl( pnrasitized grubs for shipment was the sam( I1S that described for shipment of grubs parnsitized with D ~entrais
No shipments of Prosena sibirita wer( mude aftN 1930 ns this parasite was then established in the Vnit((1 States
SCOLIIDAE
During the course of the investigations from 1920 to 1933 inclusive on the pnrasites of Popillia and related scarabaeid b(etl(s in the Orient a huge number of Scoliidne principally of the genus Tiphia were found and intensively studied ~fost of thtsP species which were consid(recl to be ntw have since been des(ribtd (2 18 19 2021)
In table 4 is presented the 1llown distribution in Tnpllll ChOS(D China Indin nncI Taiwan of all species of Tiphia real(d or shipped sinet the initilltion of the work Of the 29 sp(ei(s list(ci in this tablr 8 nre known to occur ill Japan Chosrn and China 2 in Jupnll nnd ChOS(Il and 1 in ChOS(1l and Chinn No doubt furtlH1 innstigtltions will show a much wider dissemination throughout the Orient Of the species mentioned in this bulletin only liphia bisfculata nnd T tegitiplaga purnsites of Anomala allld occurring in southern Jnpnn have 2 distinct generations a year
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
j
OQ 13
0 0 lts ~
lt ~ a ~ ~ ~ ~
gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
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L
5 IXYESTlGATIOXS OF PARASITES OF JAPAXESE BEETLE
HAMAXIA INCONGRtA Vnlklr
llam(llia inconglua a pmusite of the adults of many species of Scarabutidae is widely distributed throughout the Asiatic and IVlalaYfill Regions find in various tropicn countries In Japan where it occurs as a parasite of Popillia japonica it has two and possibly thrre genErntions a year Bping of crepuscular habitJ it is active mostly during the carlr morning hours and in the cvening Although the lxuct mlthod of parasitization is not known it is prlSUDlN that thc femult dlposits fully deClopN lalYap lither on the host beltle or on the llrnrby foliagr and that tlipse enter the beetle through thc softer portions of t111 body
Although no shipments of IJama11aincongrua have bern made to the Lnitrd StlltlS since 1928 obsllyations on distribution and degree of field pHrnsitizatioIl werc carrid on in Japan during th period 1929-33
During 1930 1931 and H)3~ onl 200000 Popillia beetlls (1(
collected nt 108 localitis lepnsenting a wide range of climatic tlnll enviroIlmeJltal conditions 011 the islnnds of Honshu Sbikoku and Kyushu in lapan These were collected from the middle of Junc to the end of August nod in some instances eol1ections were made at weekly internlls Since it is impossible to distinguish the pnrasitizpltl beetles in the field owing to the lnrvipositing habit of this tachinid all collecttd material was sent to the Yokohama laboratory for obsershyvation Hamaria incongrua was reared from beetles collected at 68 of these localiti(s on til( 3 islnnds Emelgpnee records from this collected mnterial showed that IIamaria was the moll abundant ill the hilly places cOnled with underbrush in the wnrmer lll(ns Field pamsitizatioll wns excP((lingly YftrinbJe in diffplent spdiol1s unci thl genellll 11Wrnge yely low In 110 locality were morl than 20 percent of till beetks pnrusitized and in only 13 localiti(middots WnS pnmsitizatioll a bovp 5 pCrclll t
Pnlnsi tized b(etles WlI( found ill tlll filhl in ((11 tllll lind westcIll Japan thlougholl t tll( season the highest rates of palUsitization occurring during thf latter part of June and early in July find again the latter part of August At Obuse in western Japan 51 percent oi tlle beetles were parllsitized on Jllly 9 1932 Examination of the beetlfs from the same locnlity coll(cted July 25 nnd 29 rCvealed pnrasitizutions of 23 and 32 pNcpnt respctively which incrensed to 143 percrnt of the beetlls rollcted the last week in August Popillia collected at Takao near Tokyo in enstern Japan on July 3 1932 had n parusitizfl tioD of 11 ppl(ent which dlopprd to 12 flnd 14 percen t in (ollctions made on J tlly 19 nnll 28 lesppctiVCly and rose flgain to 43 p(lcent ill a lot collected August] 9 The differencps in these palnsitization r(gtcorc18 are no doubt tlue to the two generations of llamaJmiddotia I1ppearing in thl field at tlwse times
In alN1S wlHre adults of Popillin japonicn arp prCs(nt in thC fidel over a period of ~ months or longer llam(uia in(ongrua is ab to develop two glneratiom on this host but in arcas where this dOlS not occur flltcrnatc scarnbacid host bectles a Ie lleCPSS1UT for the pnS(lshyvation of this species
6 TECHNICAL BULLETIN 738 U S DEPT OF AGRICCLTGHE
EUlRIXOPSIS J VANA Townsend
In 1923 nnd 1928 a few adults of Eutrixopsis javana were reared at the Riverton laboratory from shipmeuts of beetles parasitized by Oenteiel cinerea forwarded from Sapporo Japan in 1922 and 1927
Although several attempts haYc been made to obtain information on this tachinid in Japan nothing relative to its biology or life history has been found A cardul examination of beetles bearing tllcbinid eggs collected at Snpporo in 1928 revealed that all tlll pupuria conshytained within the dead parasitized beetles wen those of (enteter ~10re than 100000 beetles free from external tacbinid eggs were also examined but not a single Eutrixopsis puparium vas found In 1930 approximately 10000 PopilliCL weI( (oUecteci at 9 otl(r localities on Rokkaido Island ~Jore than half of the beetles bore tachinid eggs and the remainder were free from extemal eggs Luter examination of both pnrnsitizpd and unpuIfisitized beetles revealNl nothing but Centeier puparin
Apparently Eutrilopsis at times parasitizes Popillia in the absence of some other host and may be consici(led of little importance as a fnctor in the control of P j(Lponica in Inpnn
TnOlHOPS CLA lHEN Aldrich
Trophops cla1l8eni was first discoerNl in 1928 whln ~OOO Popillia iaponica brrtlps colllClrd at Tokorozawn neaL Tokyo Japan late in June rcvpaled t parasitization of 12 pCLeent Blrths (ollrcted at this arCH 1 wcek latlL hoe-el did not eontain n single individual
In 1930 while the distribution of and fi(lcl pnrnsitizltioll by (enieter cinerea find lIamaria iIlCOnflla WeIe being inv(stigatNl in XagHno prefedurr westprn Japan Troptops wns again pn(ollutrred in two locnliti(s In 1931 it wus found in sCnrnl ndditiollllllolulitips in the sumc prefpcture The highest pnlHsitiznLion WitS found at Obuse middotwherc collections of Popillia in 19~2 yielded 81 nnd i) pereeut on July 9 Hnd ] 3 rrspecti vlly The field pnrnsitization of hcptllS collected in all loealities however uytrngpd only 14 lwrccnt in that year Of seyrrnl Scarabueidae present in the fi(middotld in these localities Anomala lujocuplea appears to bc the only other beetle a(cpptrd by tlus tachinid as a host Although Popiliia heetles collpded at Toko1ozawa latp in Junc in 1928 we1P found to contnin Trophops laryae no pnrnsitiz((l beetl(s wcre found in NngnTlo prei(ctule brforc July 5 Aftel this date the degne of fipld pnrHsitizntion increnscd until the middle of the month aftrl which it grndllnlly dePlined After the first wepk in August no pnrnsitizcdPopillia could bp found although A rllocuprea coll(cted late in Allgust wpre found to rontain larvue of Trophops Colkctions of berthmiddots made in thr middle of July showed the sex ratio of the attacked brctles to be 61=3 pcrcent mnles nnd 387 perc(nl females
Votes on the L1je llistMy of Trophops claU~flti
Thp tachinid Hy Trophops clauReni is npparCntly l10eturnnl in its habits for it has never he(n obs(ry(d in Ih( filld PlrHsitizcd heetles are uSllnlly found in rather hilly pinelS with dtl1s( undelshygrowth indicating fidel habits similar to those of IIamaxia Hppellteel
7 I~TESTIGATIOXS OF PARASITES OF rAPAXESE BEETLE
examinations of the beetles containing the 1aryue of Trophops showed no traces of external rggs so it is nssuinrd that the femnle deposits living larvae but the exuct mllnnel in which it attacks its host or the manner of penetrution by the 1urva into tll( host heet1( has n(ycr been determined The clrath of the bretle occurs when the purasitc larva has reached the eurly purt of th( thin1 instal TIl( period of huva1 clevrlopmcnt in the host body is similar to that of Hamaria incongrua the ttvemgr time from roHection of the b((tles to pupation of the parasitE being 9 clays nnd(l nOlmnl t(mp(Lll tllles pr(yuiling during midsummer In July nnd August the pupnl stagr in Popiliia COWlS a prriod of 15 to 17 clnys Howrycr in Anomala r1fOC1lprea which werE collected in the latter part of August the pupal peliod was 28 clnys
The rxuct llumb(l of generations of Trophops in (8t(111 Jnpull is not known although there arr at l(ust two a year the first on Popillia und the folloing onr or morf on aailable Popillia nne Anomala lujocuprea In 1982 Tlophops en1llgrd dnring th( rnriy part of August from Popillia b(etl(s that had bern collected in the middle of July und pound1 71tfocllprea colledCd in the latter part of August were also found to contain lnrnlc of this parasite
The stntns of this tnchinid ns a pnmsite of Popillia japonica is son1lwhat similnr to that of [Jamajia incongnw although its known distribution is morE limitrd BrcHusr of its low rnJe of field pamsitishyzntion ns ohs(lwd in its natin habitat in Japan nnd its two or more gruelUtions a yrnr conring it prriod of ndult nctiity of 2 months or long(l the alur of Troplwps clallseni ns a pnrasite of P japonica in the rnitecl Stntes is questionabll
In 1932 1 totnl of 287 pupa ria of TropllOp8 clauseni erl shipPNI to the rnitNI StntrR from which only 16 adults (merged in the spring of 1933
PYHGOTlDAE
ADAPILIA FIA VIfltETA Aldrich
A full account of tIl( lif( history nnd genlral hnbits of Adapsilia jialiwta a parasite of adult Popillia blltlrs in Inclin has heen given in a previous publication (10) In Indin this pyrgotid Il~middot is restricted to the hilly sections that nre eoyered with wieldy Spltc(eI plnntings of pine trees and fnirly dll1sr undergrowth of brush nt an (elltion of about )000 flet It has only ol1r glnlration It year the ndults apppuring in thefilld about thl middll of July Its normal host in India is Popilia cllpricolli8 although P tyanea nnt P macclellandi arl occl1siol1itlly pnrllsitizrcl In parnsitizing th( bp(ll( lhr [(mnle ildapnmiddotZia which is diurnal rrmnins quirs((llt on th( foliag( nfar the feeding br(tlr until thr brrtlr tnkrs flight whr[rupon till fI~ immedishyately pOllnc(s upon thr b(rtlr from nhoH It tlwn quickly insClts its egg through [1 long rrlntiwly slend(r and shnrp ovipositor into the mid-dorsal l(gion of the b((tlr nhdom(n Il(nr the junctllle with thl thorax
Thr work in Indin in H)29 was rrvotc1rl to col1((ting hl(ltlr l1EfLl Sbillong lWei sbi pping 1dapsilia pu pnria obtai nrcl from thpsr hl(tlrs to the Cnited Stt1t(S X0 furthr work was don( with this BI)((ies nftrr 1929 owing to th( dis(ontinuntion of thr work in thttt country
Brrt1Es W(Ire collcet(l(l dllling tIl( period from July 25 to August 18
8 TECHXICAL BULLETIN 738 U S DEPT OF GRICFLTCRE
1929 th( majority being brought in from August 1 to 9 th( p(liod when the synchronization of host and parasite is at its best DlIling this time 553885 beetles we1( brought in by the llum(IOtls l1atic collectors FIOIU these 4310 Adapsilia pupalin were obtained giving a parasitization of 078 percent as compared with 053 in 1928
TIlt manuel of renring uncI shipping wus similnr to thnt (mployed in preyious yenls The beetles were confined in hla cardboard boxes of approximately 1 cubic foot capacity Illld to absorb excess moisture the bottoms w(1e lined with fresh moss Apples were used to f(ed the beetles and were l(phnisbed on alternate dllYS After 12 dnys allliye bC(tles which were (lvid(ntly Ullpn1llsitized W(Ie destroy(d nnd the dead olles placed in opm trnys to allow the Atlapsilia which W(lre still ill the Inrvn1 stngr to pupntt Artel pupntion til( UllCOIlshySUlllNl body rontrnts of th( host wrre rellloved to diminnt( decomshyposition gases that might arise durillg shipmrnt to til( detriment of the pllparia The pupnrin v(1e packed betwNm ltl~nS of fresh sphagnum moss in smull vrlltillLted 1ectnnglllnJ tillS and n number of tlllse en tirely surroUlHlpd by 1 3-illClL layer of fllsh mo (1(
pfldwl in illS(Ctploof woodell box(s for shipment One shipnwnt eonsisting of 3ROO pupnrin was SPIlt to tli( rnited
Stnt(s in 1929 This arrind at its dCstinntion Xonmb(r 9 Illvinp bN1l ell rou tp 42 ditys nt it t(mp(rntue of 40deg to 50 0 F
PARASITES OF TilE LAIVA
nEXIlDAE
DEXIA EXTnTIS Aldrich
Delin 1flltrali tilr most rommon of the ChmltHs( dlxiids Wiltshyfirst found Ilt SUiglll ChosllI in 1922 (9) Owing to its (olllplirntld biology no r(Hring work for the shipnwnt of pnmsitiz(d glU bs to t1H lnit(gtd StatN was aWmpted until 1)21) (10) In that Yenr P()pilli(t jnponic(L grubs whirh do not orcur in Chosen (1( eol1letrd filld shipped from nortlH11l Japan to Fukuoka in Kyusitu soutitNll Jnpitn for pnrnsitizlltion by Duia (mnks fonnumiddottkd from Chos) By thr tinlP n suiJici(nt numb(middot of glflYid f(BmTes Wtl( nvailablr Inte i11 JU11r how(yrr most of the grubs w(re too far d(vdop(d for pnrnshysitiztltion purPOR(S find only 850 pnrnsitizld grubs were shipped to til( enitrd States
In 1926 it WfiS d((idpd to (nlTY on rlaring work in 0hos1 utilizing the nutiY( ChOSlll(RC host grubs This provrd to be very sucressful and OWl 20000 grubs were pamsitizrd flnd shippld to titC rnit((l States in 102(3 lind 1927 B(ctlUSC oJ til( dlLngPl iIlYoiwcl in shipping to the rnited Statts Jive grubs othrl thtlll tilos( of POJlillia iapflnica owing to tIl( risk of 111 rscnpl of thrse sp(cils no l(uring w(lIk WIlf
cHrrild on in Chosen in 1928 but 11 mdhod WIIS dlvCloppd whplpby adult fCmale DIJia could be sitipPllaquo LO Yokollilma SII(clssfully and in 1029 rrnring work wns rnrri(d on til(lC the grll bs of P japof ira being IIs((I tiS hosts
Tabl( 2 shows the slllTinl of ndult f(malCs of [Jain middot(lltr(1Ii forshywllrd((l from SlIig(Il Chospn to YokohfilJ11L in 1029 Hnl 1930 til( yciUS that rCaring work WilS el1JTi(d OIl TIle I1dults PI( ronfillNI in thl shipping containl1s which WCI( similm to thos llIlployltd ill transporting Scoliidne npprox11llat(ly 7 days from the timr o( (oll(eshytion to the tim( of removaL
IXVESTIGATIOXS OF PARASITES OF TAPANK~ BEETLE 9
TABLE 2-SuTviLal of adult females of DeJia ventralis shipped from 81tigcn Chosen 10 Yokohama Japa11in 1929 and 1930
Adults Adults surllingYear Generntion shipped shipment
Numb lrumber PercentI
1929_ bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull _ FirsLbullbullbullbullbullbullbullbullbullbullbullbull _bull __ bull __ bullbull_1 063 552 833 1921 __ bullbullbullbullbull ______ bull __ bullbull ____ lhird____ __ 2281 1800 831 1930 __ _bullbullbullbullbullbull __ bull __ First _______ __ ~9() 6S- 68 1910_ _ __ _____ bull_____ __ 1 erond_____ __ __ 2051 14il 720 1930bullbullbull __ bullbullbullbull __ Third __ 2483 2008 809
Total __ __ __ ___ 8368 t 661 ]----()-1 I
It was extremrly difficult to obtain Popillia grubs in any numbers in the --icinity of Yokohamn for rearing work with Deria but this was OYerC0111( by collecting ancl shipping grubs as IHeclNI from Hokshykaido in nortlltrn Japall to Yokohallla n procedure which proyrd to be yery satisfactory and inrxplnsiYe ~0 rearing work was attempted with the s(cond gel1lrntion of DeJia in 1929 oing to the impossibility of s(curing u sufficient number of grubs in IIokkuido during August This was the YNlr of grub scarcity in northern Japall vhen less than 1 percent of the oYCrwintering grubs huying a 2-y(middotnr lifc cyclc (le still in the brynl stage at thn t time of the yeal
The ptlrasitizNl grubs obtain(d tluough parasitization with the first and s(concl g(l1Plntions of DU(L in June nnd August lPspCctiYely wele shipped to thpCnitltC1 Stat(s immpdiatrly aftC1 btinf pnra~itizcd DcYClopment of the parasites took pla(e P11 route nnd the adults emrTgecl sho1tl)- uiter nrriyal at tlHlT destination Grubs containing tbe hibernnting ln1yue of De ia obtuined through pnrasitizatiOIl with the third-g(lHlntion 1Ii(s in 8(ptp111her nnd Octol)(r Pre kPpt OY(1 wintpr at 10kohi1111tl in n wlll-crain(c plot (oY(INI with sod The winttr lllortnlity of these grubs was rnthel high ns only 171 and HL9 pcrcent werc nIl( when dug up [or shipnlPnt in the springs of 1930 lllHl 1931 rlspertiwly
Rwring and Shipping Jfpoundt7wds
The rcuring work with lJJ(l l(fltl(llj1 was accomplisll(d hy using a bnmiddotpding-jnl 111Cthod simillll to that employrd in ]Jle-iolls y(nrs Approximately 50 grubs (Ie placfll in the lonrhalf of a l-qualmiddott glnss jar with soil A fin(-IJ1Psh SCIepn cngp contllining 5 or () lJ(tia f(malp WIlS fittNI into this jur lind rPRtpd on til( soil nnd through this scnPI1 into the soil til( dlpositcd lalTHP pnsNI in palch of the host gruhs 1llP grubs wpl(~ (xtunilHd pneh chty Hlld thORP showing thf pnrnsit( nmiddotpjlntol~ funlwl (1( ((monel nne Tplacpd with frlsh g-rub The flips which Pl( fpd IPg III ll[l with it solution of sunr und watN linmiddotd f[om 3 middot(middotCk tc) J IJ1fll1th in til( cage nnd pnrnitizld flom a5 to )0 pP[cpnt of tlH grub Iach dnv
In pucking the pnrnsitizplt1 gruh 101 shipnwnt iin-lillpd hoxls 1 foot squnrp und Ii inch(s dNp W(IP wwd Each l)()x contnilpd 5 cro-5lction s(ts RPparutd b lin pintp ploviding it COll1pnltllHnt of nhout 1 (lbic inch foJ Pilch of jOIl glubs Fol shipllllllt 2 of thpse hoxps (1( packlrl togtlwl in n (Tong woodpl1 (nsp TIl( soil in which thp gru bs (n pu(k(d had hren thorough Iy fumign tld wi t h (arbon (hsulfid( to (limirmtC any otlw1 inlcl lllntrrial which might htwe
23-13-1----40-2
]0 TECH~ICAL BULLETIN 738 U S DEPT OF AGRICULTURE
been present In table 3 is a summary of shipments of palasitized grubs and the percentages of parasite emcrgenee after arrivul in thl United Stntls
TABLE 3-SlImmary oj shipments oj Popillia japonica grubs l)ara~itizcd willi Deda Ientralis Jrom Japan to the United States 1929-31
Year Contents oC shipment Grubs Parnsites emergingship(led-------1--------------- ---- ______ lVumber lVlLmber I Pentnt
19211-bullbullbullbullbullbullbullbullbullbull_bullbull Grubs pnmsiti1ed with flrstgenerntion Dezia ~ ~500 ~51 1000 1930_bullbullbull __ bull __ bullbullbullbullbull __ Overwintered grubs parnsitized with thirdmiddot 3--
gcnemtionDaiu ~~_~ __ ~~ ___ _ ___ ~ 3000 II 126 1930bullbullbull ___bullbullbullbullbull __bull ___ Grubs pamsitized with lirst-enerntion Detia~~ 8000 1504 18S 1930_____________ ~ _ __ Orubs pllrasitized with ~econd-generntion Drtia 7000 l42J I 203 1930____ __ ~ ~ ~ _ GrUbs pllmsiti1ed with second-genemtion 2500 399 16 (f
Dufu II1931___________ Oerwintered grubs paraSitized with third- 4000 345 8 ( generation Duia I
I 1--------- shyTotaJorneruge~_i- ___ bullbullbullbull __ _ _______ bull ___ ~ ~ ~)(JOOO I 4499middot 155 f i i
Owing to the establishment of Diria lentrali~ in tIl( United Stutes from which materinl for lccolonizlltion was availabll no shipments of this pfllflsitl Wlre made aftlr 1931
PnOSENA SIBII1TA (Fabricins)
Thl dexiid fly Prosena sibi7ita middotliich IS glllclfllly distrihut(d throughout till Asiatic and ~IalaYlln Hegions is an importnnt pilll1shy
sill of PopWia iaponica in Kolth(1l1 Japan It hns olle glllelation a YfIU ami is tllpubl( of palllsitizing and dfwloping Oll numplou sppcils of sealt1bnpid grubs of suitnblr size The life history nnd ililbits of this dexiicl huve been fullv dPidt with in previolls publieashytions (9 10) bull
The shipments from 1923 to H)27 inclusi-e cOl1[-iistN[ of fitldshycolkrtfd grubs of Popillia iap(mit(l whieh showld a getHrnI fitld parflsitization of 10 to 15 percpnt Becaust of till impossibility of detCrting the plesenee of the pUlllsite latnt whieh is nttuchNI to tlH main trnelwal trunk of the host all collpcted glUbs W(I( shipp((L Tllf grubs (re dug in Hoklmido rudy in the spring lind imnHdilltely forwarded to the rnited States arriving thete shortly beforp tll( emlIgenee of the parasites Although this method proed sntisshyfactory the trllnsportntioll costs wPle (xtrCll1(ly high nnd in 1927 it was dfcided to palllsitize till grubs nrtifjeiall)- by 11 mltthod that hnd bCll1 laquo(eloped in 1921 This ms aecomplished by dissecting til( first-stag( larval from tIle oVlsacs of the glUyid femuks and trilllSshy
ferring them to thC host grubs by mcans of a fine blllsh To instlll penetration by the parnsites the grubs with til( smal lnlYllc wIe vlaeed in cclIs of suitable size in wooden bloek fotmiddot it petiot of 24 hours brron being plaecd in thc field whftP tllr oYClwintrlld
Late in the summer of 1928 40596 grubs W(Il artificially PUIllshysitized in 1l0rthCrTJ Japan llnd placed in a prcpnred soil plot w(11 drained coered with sod and surrounded to a depth of 18 inch(s by a fine-mesh screen to pr(vent the gnlbs from wumlcring nwny T11(s( grubs were dug up carly in May 1929 and 19440 were nliv( of whieh 19000 weIpound shipped to the U ni ted Stat(s
In 1929 it was impossibl( to seure sufficient grubs for inoeulatioll work Consequently it was necessary to colleet Prosena femulCs and
IXETrGATIOXS OF PARASITES 01 rAlANFJSFJ DEETLB 11
ship them to the Yokohnma laboTfitory und carryon palasitization work there with available current-year grubs from that vicinity Only 12 p(rcent of the females were alive on arrival at Yokohama Tlus high mortality WitS due to the extremely hot weather prevailing at the time of shipment Out of 364 females received alive 159 contained Inrvae sufficiently developed for this work During the latter part of August 6300 grubs were parasitized and placed in hibernation qunrters As e~perienced with the overwinter~d grubs parasitized with Dexia 1entralis the wintPI mortality was (xceedingly high in this material and only 900 were alive wh(n dug up the followshying spring These with the 19000 shipped in 1929 together rith the 29000 of DeJia tentralis make a total of 48900 grubs containing dexiids thnt were shipped to the United States during the period covcl(d by this bulletin The method of pl1cking tIl( pnrasitized grubs for shipment was the sam( I1S that described for shipment of grubs parnsitized with D ~entrais
No shipments of Prosena sibirita wer( mude aftN 1930 ns this parasite was then established in the Vnit((1 States
SCOLIIDAE
During the course of the investigations from 1920 to 1933 inclusive on the pnrasites of Popillia and related scarabaeid b(etl(s in the Orient a huge number of Scoliidne principally of the genus Tiphia were found and intensively studied ~fost of thtsP species which were consid(recl to be ntw have since been des(ribtd (2 18 19 2021)
In table 4 is presented the 1llown distribution in Tnpllll ChOS(D China Indin nncI Taiwan of all species of Tiphia real(d or shipped sinet the initilltion of the work Of the 29 sp(ei(s list(ci in this tablr 8 nre known to occur ill Japan Chosrn and China 2 in Jupnll nnd ChOS(Il and 1 in ChOS(1l and Chinn No doubt furtlH1 innstigtltions will show a much wider dissemination throughout the Orient Of the species mentioned in this bulletin only liphia bisfculata nnd T tegitiplaga purnsites of Anomala allld occurring in southern Jnpnn have 2 distinct generations a year
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
j
OQ 13
0 0 lts ~
lt ~ a ~ ~ ~ ~
gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
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_ - shy__ middot_rmiddot
is
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L
6 TECHNICAL BULLETIN 738 U S DEPT OF AGRICCLTGHE
EUlRIXOPSIS J VANA Townsend
In 1923 nnd 1928 a few adults of Eutrixopsis javana were reared at the Riverton laboratory from shipmeuts of beetles parasitized by Oenteiel cinerea forwarded from Sapporo Japan in 1922 and 1927
Although several attempts haYc been made to obtain information on this tachinid in Japan nothing relative to its biology or life history has been found A cardul examination of beetles bearing tllcbinid eggs collected at Snpporo in 1928 revealed that all tlll pupuria conshytained within the dead parasitized beetles wen those of (enteter ~10re than 100000 beetles free from external tacbinid eggs were also examined but not a single Eutrixopsis puparium vas found In 1930 approximately 10000 PopilliCL weI( (oUecteci at 9 otl(r localities on Rokkaido Island ~Jore than half of the beetles bore tachinid eggs and the remainder were free from extemal eggs Luter examination of both pnrnsitizpd and unpuIfisitized beetles revealNl nothing but Centeier puparin
Apparently Eutrilopsis at times parasitizes Popillia in the absence of some other host and may be consici(led of little importance as a fnctor in the control of P j(Lponica in Inpnn
TnOlHOPS CLA lHEN Aldrich
Trophops cla1l8eni was first discoerNl in 1928 whln ~OOO Popillia iaponica brrtlps colllClrd at Tokorozawn neaL Tokyo Japan late in June rcvpaled t parasitization of 12 pCLeent Blrths (ollrcted at this arCH 1 wcek latlL hoe-el did not eontain n single individual
In 1930 while the distribution of and fi(lcl pnrnsitizltioll by (enieter cinerea find lIamaria iIlCOnflla WeIe being inv(stigatNl in XagHno prefedurr westprn Japan Troptops wns again pn(ollutrred in two locnliti(s In 1931 it wus found in sCnrnl ndditiollllllolulitips in the sumc prefpcture The highest pnlHsitiznLion WitS found at Obuse middotwherc collections of Popillia in 19~2 yielded 81 nnd i) pereeut on July 9 Hnd ] 3 rrspecti vlly The field pnrnsitization of hcptllS collected in all loealities however uytrngpd only 14 lwrccnt in that year Of seyrrnl Scarabueidae present in the fi(middotld in these localities Anomala lujocuplea appears to bc the only other beetle a(cpptrd by tlus tachinid as a host Although Popiliia heetles collpded at Toko1ozawa latp in Junc in 1928 we1P found to contnin Trophops laryae no pnrnsitiz((l beetl(s wcre found in NngnTlo prei(ctule brforc July 5 Aftel this date the degne of fipld pnrHsitizntion increnscd until the middle of the month aftrl which it grndllnlly dePlined After the first wepk in August no pnrnsitizcdPopillia could bp found although A rllocuprea coll(cted late in Allgust wpre found to rontain larvue of Trophops Colkctions of berthmiddots made in thr middle of July showed the sex ratio of the attacked brctles to be 61=3 pcrcent mnles nnd 387 perc(nl females
Votes on the L1je llistMy of Trophops claU~flti
Thp tachinid Hy Trophops clauReni is npparCntly l10eturnnl in its habits for it has never he(n obs(ry(d in Ih( filld PlrHsitizcd heetles are uSllnlly found in rather hilly pinelS with dtl1s( undelshygrowth indicating fidel habits similar to those of IIamaxia Hppellteel
7 I~TESTIGATIOXS OF PARASITES OF rAPAXESE BEETLE
examinations of the beetles containing the 1aryue of Trophops showed no traces of external rggs so it is nssuinrd that the femnle deposits living larvae but the exuct mllnnel in which it attacks its host or the manner of penetrution by the 1urva into tll( host heet1( has n(ycr been determined The clrath of the bretle occurs when the purasitc larva has reached the eurly purt of th( thin1 instal TIl( period of huva1 clevrlopmcnt in the host body is similar to that of Hamaria incongrua the ttvemgr time from roHection of the b((tles to pupation of the parasitE being 9 clays nnd(l nOlmnl t(mp(Lll tllles pr(yuiling during midsummer In July nnd August the pupnl stagr in Popiliia COWlS a prriod of 15 to 17 clnys Howrycr in Anomala r1fOC1lprea which werE collected in the latter part of August the pupal peliod was 28 clnys
The rxuct llumb(l of generations of Trophops in (8t(111 Jnpull is not known although there arr at l(ust two a year the first on Popillia und the folloing onr or morf on aailable Popillia nne Anomala lujocuprea In 1982 Tlophops en1llgrd dnring th( rnriy part of August from Popillia b(etl(s that had bern collected in the middle of July und pound1 71tfocllprea colledCd in the latter part of August were also found to contain lnrnlc of this parasite
The stntns of this tnchinid ns a pnmsite of Popillia japonica is son1lwhat similnr to that of [Jamajia incongnw although its known distribution is morE limitrd BrcHusr of its low rnJe of field pamsitishyzntion ns ohs(lwd in its natin habitat in Japan nnd its two or more gruelUtions a yrnr conring it prriod of ndult nctiity of 2 months or long(l the alur of Troplwps clallseni ns a pnrasite of P japonica in the rnitecl Stntes is questionabll
In 1932 1 totnl of 287 pupa ria of TropllOp8 clauseni erl shipPNI to the rnitNI StntrR from which only 16 adults (merged in the spring of 1933
PYHGOTlDAE
ADAPILIA FIA VIfltETA Aldrich
A full account of tIl( lif( history nnd genlral hnbits of Adapsilia jialiwta a parasite of adult Popillia blltlrs in Inclin has heen given in a previous publication (10) In Indin this pyrgotid Il~middot is restricted to the hilly sections that nre eoyered with wieldy Spltc(eI plnntings of pine trees and fnirly dll1sr undergrowth of brush nt an (elltion of about )000 flet It has only ol1r glnlration It year the ndults apppuring in thefilld about thl middll of July Its normal host in India is Popilia cllpricolli8 although P tyanea nnt P macclellandi arl occl1siol1itlly pnrllsitizrcl In parnsitizing th( bp(ll( lhr [(mnle ildapnmiddotZia which is diurnal rrmnins quirs((llt on th( foliag( nfar the feeding br(tlr until thr brrtlr tnkrs flight whr[rupon till fI~ immedishyately pOllnc(s upon thr b(rtlr from nhoH It tlwn quickly insClts its egg through [1 long rrlntiwly slend(r and shnrp ovipositor into the mid-dorsal l(gion of the b((tlr nhdom(n Il(nr the junctllle with thl thorax
Thr work in Indin in H)29 was rrvotc1rl to col1((ting hl(ltlr l1EfLl Sbillong lWei sbi pping 1dapsilia pu pnria obtai nrcl from thpsr hl(tlrs to the Cnited Stt1t(S X0 furthr work was don( with this BI)((ies nftrr 1929 owing to th( dis(ontinuntion of thr work in thttt country
Brrt1Es W(Ire collcet(l(l dllling tIl( period from July 25 to August 18
8 TECHXICAL BULLETIN 738 U S DEPT OF GRICFLTCRE
1929 th( majority being brought in from August 1 to 9 th( p(liod when the synchronization of host and parasite is at its best DlIling this time 553885 beetles we1( brought in by the llum(IOtls l1atic collectors FIOIU these 4310 Adapsilia pupalin were obtained giving a parasitization of 078 percent as compared with 053 in 1928
TIlt manuel of renring uncI shipping wus similnr to thnt (mployed in preyious yenls The beetles were confined in hla cardboard boxes of approximately 1 cubic foot capacity Illld to absorb excess moisture the bottoms w(1e lined with fresh moss Apples were used to f(ed the beetles and were l(phnisbed on alternate dllYS After 12 dnys allliye bC(tles which were (lvid(ntly Ullpn1llsitized W(Ie destroy(d nnd the dead olles placed in opm trnys to allow the Atlapsilia which W(lre still ill the Inrvn1 stngr to pupntt Artel pupntion til( UllCOIlshySUlllNl body rontrnts of th( host wrre rellloved to diminnt( decomshyposition gases that might arise durillg shipmrnt to til( detriment of the pllparia The pupnrin v(1e packed betwNm ltl~nS of fresh sphagnum moss in smull vrlltillLted 1ectnnglllnJ tillS and n number of tlllse en tirely surroUlHlpd by 1 3-illClL layer of fllsh mo (1(
pfldwl in illS(Ctploof woodell box(s for shipment One shipnwnt eonsisting of 3ROO pupnrin was SPIlt to tli( rnited
Stnt(s in 1929 This arrind at its dCstinntion Xonmb(r 9 Illvinp bN1l ell rou tp 42 ditys nt it t(mp(rntue of 40deg to 50 0 F
PARASITES OF TilE LAIVA
nEXIlDAE
DEXIA EXTnTIS Aldrich
Delin 1flltrali tilr most rommon of the ChmltHs( dlxiids Wiltshyfirst found Ilt SUiglll ChosllI in 1922 (9) Owing to its (olllplirntld biology no r(Hring work for the shipnwnt of pnmsitiz(d glU bs to t1H lnit(gtd StatN was aWmpted until 1)21) (10) In that Yenr P()pilli(t jnponic(L grubs whirh do not orcur in Chosen (1( eol1letrd filld shipped from nortlH11l Japan to Fukuoka in Kyusitu soutitNll Jnpitn for pnrnsitizlltion by Duia (mnks fonnumiddottkd from Chos) By thr tinlP n suiJici(nt numb(middot of glflYid f(BmTes Wtl( nvailablr Inte i11 JU11r how(yrr most of the grubs w(re too far d(vdop(d for pnrnshysitiztltion purPOR(S find only 850 pnrnsitizld grubs were shipped to til( enitrd States
In 1926 it WfiS d((idpd to (nlTY on rlaring work in 0hos1 utilizing the nutiY( ChOSlll(RC host grubs This provrd to be very sucressful and OWl 20000 grubs were pamsitizrd flnd shippld to titC rnit((l States in 102(3 lind 1927 B(ctlUSC oJ til( dlLngPl iIlYoiwcl in shipping to the rnited Statts Jive grubs othrl thtlll tilos( of POJlillia iapflnica owing to tIl( risk of 111 rscnpl of thrse sp(cils no l(uring w(lIk WIlf
cHrrild on in Chosen in 1928 but 11 mdhod WIIS dlvCloppd whplpby adult fCmale DIJia could be sitipPllaquo LO Yokollilma SII(clssfully and in 1029 rrnring work wns rnrri(d on til(lC the grll bs of P japof ira being IIs((I tiS hosts
Tabl( 2 shows the slllTinl of ndult f(malCs of [Jain middot(lltr(1Ii forshywllrd((l from SlIig(Il Chospn to YokohfilJ11L in 1029 Hnl 1930 til( yciUS that rCaring work WilS el1JTi(d OIl TIle I1dults PI( ronfillNI in thl shipping containl1s which WCI( similm to thos llIlployltd ill transporting Scoliidne npprox11llat(ly 7 days from the timr o( (oll(eshytion to the tim( of removaL
IXVESTIGATIOXS OF PARASITES OF TAPANK~ BEETLE 9
TABLE 2-SuTviLal of adult females of DeJia ventralis shipped from 81tigcn Chosen 10 Yokohama Japa11in 1929 and 1930
Adults Adults surllingYear Generntion shipped shipment
Numb lrumber PercentI
1929_ bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull _ FirsLbullbullbullbullbullbullbullbullbullbullbullbull _bull __ bull __ bullbull_1 063 552 833 1921 __ bullbullbullbullbull ______ bull __ bullbull ____ lhird____ __ 2281 1800 831 1930 __ _bullbullbullbullbullbull __ bull __ First _______ __ ~9() 6S- 68 1910_ _ __ _____ bull_____ __ 1 erond_____ __ __ 2051 14il 720 1930bullbullbull __ bullbullbullbull __ Third __ 2483 2008 809
Total __ __ __ ___ 8368 t 661 ]----()-1 I
It was extremrly difficult to obtain Popillia grubs in any numbers in the --icinity of Yokohamn for rearing work with Deria but this was OYerC0111( by collecting ancl shipping grubs as IHeclNI from Hokshykaido in nortlltrn Japall to Yokohallla n procedure which proyrd to be yery satisfactory and inrxplnsiYe ~0 rearing work was attempted with the s(cond gel1lrntion of DeJia in 1929 oing to the impossibility of s(curing u sufficient number of grubs in IIokkuido during August This was the YNlr of grub scarcity in northern Japall vhen less than 1 percent of the oYCrwintering grubs huying a 2-y(middotnr lifc cyclc (le still in the brynl stage at thn t time of the yeal
The ptlrasitizNl grubs obtain(d tluough parasitization with the first and s(concl g(l1Plntions of DU(L in June nnd August lPspCctiYely wele shipped to thpCnitltC1 Stat(s immpdiatrly aftC1 btinf pnra~itizcd DcYClopment of the parasites took pla(e P11 route nnd the adults emrTgecl sho1tl)- uiter nrriyal at tlHlT destination Grubs containing tbe hibernnting ln1yue of De ia obtuined through pnrasitizatiOIl with the third-g(lHlntion 1Ii(s in 8(ptp111her nnd Octol)(r Pre kPpt OY(1 wintpr at 10kohi1111tl in n wlll-crain(c plot (oY(INI with sod The winttr lllortnlity of these grubs was rnthel high ns only 171 and HL9 pcrcent werc nIl( when dug up [or shipnlPnt in the springs of 1930 lllHl 1931 rlspertiwly
Rwring and Shipping Jfpoundt7wds
The rcuring work with lJJ(l l(fltl(llj1 was accomplisll(d hy using a bnmiddotpding-jnl 111Cthod simillll to that employrd in ]Jle-iolls y(nrs Approximately 50 grubs (Ie placfll in the lonrhalf of a l-qualmiddott glnss jar with soil A fin(-IJ1Psh SCIepn cngp contllining 5 or () lJ(tia f(malp WIlS fittNI into this jur lind rPRtpd on til( soil nnd through this scnPI1 into the soil til( dlpositcd lalTHP pnsNI in palch of the host gruhs 1llP grubs wpl(~ (xtunilHd pneh chty Hlld thORP showing thf pnrnsit( nmiddotpjlntol~ funlwl (1( ((monel nne Tplacpd with frlsh g-rub The flips which Pl( fpd IPg III ll[l with it solution of sunr und watN linmiddotd f[om 3 middot(middotCk tc) J IJ1fll1th in til( cage nnd pnrnitizld flom a5 to )0 pP[cpnt of tlH grub Iach dnv
In pucking the pnrnsitizplt1 gruh 101 shipnwnt iin-lillpd hoxls 1 foot squnrp und Ii inch(s dNp W(IP wwd Each l)()x contnilpd 5 cro-5lction s(ts RPparutd b lin pintp ploviding it COll1pnltllHnt of nhout 1 (lbic inch foJ Pilch of jOIl glubs Fol shipllllllt 2 of thpse hoxps (1( packlrl togtlwl in n (Tong woodpl1 (nsp TIl( soil in which thp gru bs (n pu(k(d had hren thorough Iy fumign tld wi t h (arbon (hsulfid( to (limirmtC any otlw1 inlcl lllntrrial which might htwe
23-13-1----40-2
]0 TECH~ICAL BULLETIN 738 U S DEPT OF AGRICULTURE
been present In table 3 is a summary of shipments of palasitized grubs and the percentages of parasite emcrgenee after arrivul in thl United Stntls
TABLE 3-SlImmary oj shipments oj Popillia japonica grubs l)ara~itizcd willi Deda Ientralis Jrom Japan to the United States 1929-31
Year Contents oC shipment Grubs Parnsites emergingship(led-------1--------------- ---- ______ lVumber lVlLmber I Pentnt
19211-bullbullbullbullbullbullbullbullbullbull_bullbull Grubs pnmsiti1ed with flrstgenerntion Dezia ~ ~500 ~51 1000 1930_bullbullbull __ bull __ bullbullbullbullbull __ Overwintered grubs parnsitized with thirdmiddot 3--
gcnemtionDaiu ~~_~ __ ~~ ___ _ ___ ~ 3000 II 126 1930bullbullbull ___bullbullbullbullbull __bull ___ Grubs pamsitized with lirst-enerntion Detia~~ 8000 1504 18S 1930_____________ ~ _ __ Orubs pllrasitized with ~econd-generntion Drtia 7000 l42J I 203 1930____ __ ~ ~ ~ _ GrUbs pllmsiti1ed with second-genemtion 2500 399 16 (f
Dufu II1931___________ Oerwintered grubs paraSitized with third- 4000 345 8 ( generation Duia I
I 1--------- shyTotaJorneruge~_i- ___ bullbullbullbull __ _ _______ bull ___ ~ ~ ~)(JOOO I 4499middot 155 f i i
Owing to the establishment of Diria lentrali~ in tIl( United Stutes from which materinl for lccolonizlltion was availabll no shipments of this pfllflsitl Wlre made aftlr 1931
PnOSENA SIBII1TA (Fabricins)
Thl dexiid fly Prosena sibi7ita middotliich IS glllclfllly distrihut(d throughout till Asiatic and ~IalaYlln Hegions is an importnnt pilll1shy
sill of PopWia iaponica in Kolth(1l1 Japan It hns olle glllelation a YfIU ami is tllpubl( of palllsitizing and dfwloping Oll numplou sppcils of sealt1bnpid grubs of suitnblr size The life history nnd ililbits of this dexiicl huve been fullv dPidt with in previolls publieashytions (9 10) bull
The shipments from 1923 to H)27 inclusi-e cOl1[-iistN[ of fitldshycolkrtfd grubs of Popillia iap(mit(l whieh showld a getHrnI fitld parflsitization of 10 to 15 percpnt Becaust of till impossibility of detCrting the plesenee of the pUlllsite latnt whieh is nttuchNI to tlH main trnelwal trunk of the host all collpcted glUbs W(I( shipp((L Tllf grubs (re dug in Hoklmido rudy in the spring lind imnHdilltely forwarded to the rnited States arriving thete shortly beforp tll( emlIgenee of the parasites Although this method proed sntisshyfactory the trllnsportntioll costs wPle (xtrCll1(ly high nnd in 1927 it was dfcided to palllsitize till grubs nrtifjeiall)- by 11 mltthod that hnd bCll1 laquo(eloped in 1921 This ms aecomplished by dissecting til( first-stag( larval from tIle oVlsacs of the glUyid femuks and trilllSshy
ferring them to thC host grubs by mcans of a fine blllsh To instlll penetration by the parnsites the grubs with til( smal lnlYllc wIe vlaeed in cclIs of suitable size in wooden bloek fotmiddot it petiot of 24 hours brron being plaecd in thc field whftP tllr oYClwintrlld
Late in the summer of 1928 40596 grubs W(Il artificially PUIllshysitized in 1l0rthCrTJ Japan llnd placed in a prcpnred soil plot w(11 drained coered with sod and surrounded to a depth of 18 inch(s by a fine-mesh screen to pr(vent the gnlbs from wumlcring nwny T11(s( grubs were dug up carly in May 1929 and 19440 were nliv( of whieh 19000 weIpound shipped to the U ni ted Stat(s
In 1929 it was impossibl( to seure sufficient grubs for inoeulatioll work Consequently it was necessary to colleet Prosena femulCs and
IXETrGATIOXS OF PARASITES 01 rAlANFJSFJ DEETLB 11
ship them to the Yokohnma laboTfitory und carryon palasitization work there with available current-year grubs from that vicinity Only 12 p(rcent of the females were alive on arrival at Yokohama Tlus high mortality WitS due to the extremely hot weather prevailing at the time of shipment Out of 364 females received alive 159 contained Inrvae sufficiently developed for this work During the latter part of August 6300 grubs were parasitized and placed in hibernation qunrters As e~perienced with the overwinter~d grubs parasitized with Dexia 1entralis the wintPI mortality was (xceedingly high in this material and only 900 were alive wh(n dug up the followshying spring These with the 19000 shipped in 1929 together rith the 29000 of DeJia tentralis make a total of 48900 grubs containing dexiids thnt were shipped to the United States during the period covcl(d by this bulletin The method of pl1cking tIl( pnrasitized grubs for shipment was the sam( I1S that described for shipment of grubs parnsitized with D ~entrais
No shipments of Prosena sibirita wer( mude aftN 1930 ns this parasite was then established in the Vnit((1 States
SCOLIIDAE
During the course of the investigations from 1920 to 1933 inclusive on the pnrasites of Popillia and related scarabaeid b(etl(s in the Orient a huge number of Scoliidne principally of the genus Tiphia were found and intensively studied ~fost of thtsP species which were consid(recl to be ntw have since been des(ribtd (2 18 19 2021)
In table 4 is presented the 1llown distribution in Tnpllll ChOS(D China Indin nncI Taiwan of all species of Tiphia real(d or shipped sinet the initilltion of the work Of the 29 sp(ei(s list(ci in this tablr 8 nre known to occur ill Japan Chosrn and China 2 in Jupnll nnd ChOS(Il and 1 in ChOS(1l and Chinn No doubt furtlH1 innstigtltions will show a much wider dissemination throughout the Orient Of the species mentioned in this bulletin only liphia bisfculata nnd T tegitiplaga purnsites of Anomala allld occurring in southern Jnpnn have 2 distinct generations a year
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
j
OQ 13
0 0 lts ~
lt ~ a ~ ~ ~ ~
gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
7 I~TESTIGATIOXS OF PARASITES OF rAPAXESE BEETLE
examinations of the beetles containing the 1aryue of Trophops showed no traces of external rggs so it is nssuinrd that the femnle deposits living larvae but the exuct mllnnel in which it attacks its host or the manner of penetrution by the 1urva into tll( host heet1( has n(ycr been determined The clrath of the bretle occurs when the purasitc larva has reached the eurly purt of th( thin1 instal TIl( period of huva1 clevrlopmcnt in the host body is similar to that of Hamaria incongrua the ttvemgr time from roHection of the b((tles to pupation of the parasitE being 9 clays nnd(l nOlmnl t(mp(Lll tllles pr(yuiling during midsummer In July nnd August the pupnl stagr in Popiliia COWlS a prriod of 15 to 17 clnys Howrycr in Anomala r1fOC1lprea which werE collected in the latter part of August the pupal peliod was 28 clnys
The rxuct llumb(l of generations of Trophops in (8t(111 Jnpull is not known although there arr at l(ust two a year the first on Popillia und the folloing onr or morf on aailable Popillia nne Anomala lujocuprea In 1982 Tlophops en1llgrd dnring th( rnriy part of August from Popillia b(etl(s that had bern collected in the middle of July und pound1 71tfocllprea colledCd in the latter part of August were also found to contain lnrnlc of this parasite
The stntns of this tnchinid ns a pnmsite of Popillia japonica is son1lwhat similnr to that of [Jamajia incongnw although its known distribution is morE limitrd BrcHusr of its low rnJe of field pamsitishyzntion ns ohs(lwd in its natin habitat in Japan nnd its two or more gruelUtions a yrnr conring it prriod of ndult nctiity of 2 months or long(l the alur of Troplwps clallseni ns a pnrasite of P japonica in the rnitecl Stntes is questionabll
In 1932 1 totnl of 287 pupa ria of TropllOp8 clauseni erl shipPNI to the rnitNI StntrR from which only 16 adults (merged in the spring of 1933
PYHGOTlDAE
ADAPILIA FIA VIfltETA Aldrich
A full account of tIl( lif( history nnd genlral hnbits of Adapsilia jialiwta a parasite of adult Popillia blltlrs in Inclin has heen given in a previous publication (10) In Indin this pyrgotid Il~middot is restricted to the hilly sections that nre eoyered with wieldy Spltc(eI plnntings of pine trees and fnirly dll1sr undergrowth of brush nt an (elltion of about )000 flet It has only ol1r glnlration It year the ndults apppuring in thefilld about thl middll of July Its normal host in India is Popilia cllpricolli8 although P tyanea nnt P macclellandi arl occl1siol1itlly pnrllsitizrcl In parnsitizing th( bp(ll( lhr [(mnle ildapnmiddotZia which is diurnal rrmnins quirs((llt on th( foliag( nfar the feeding br(tlr until thr brrtlr tnkrs flight whr[rupon till fI~ immedishyately pOllnc(s upon thr b(rtlr from nhoH It tlwn quickly insClts its egg through [1 long rrlntiwly slend(r and shnrp ovipositor into the mid-dorsal l(gion of the b((tlr nhdom(n Il(nr the junctllle with thl thorax
Thr work in Indin in H)29 was rrvotc1rl to col1((ting hl(ltlr l1EfLl Sbillong lWei sbi pping 1dapsilia pu pnria obtai nrcl from thpsr hl(tlrs to the Cnited Stt1t(S X0 furthr work was don( with this BI)((ies nftrr 1929 owing to th( dis(ontinuntion of thr work in thttt country
Brrt1Es W(Ire collcet(l(l dllling tIl( period from July 25 to August 18
8 TECHXICAL BULLETIN 738 U S DEPT OF GRICFLTCRE
1929 th( majority being brought in from August 1 to 9 th( p(liod when the synchronization of host and parasite is at its best DlIling this time 553885 beetles we1( brought in by the llum(IOtls l1atic collectors FIOIU these 4310 Adapsilia pupalin were obtained giving a parasitization of 078 percent as compared with 053 in 1928
TIlt manuel of renring uncI shipping wus similnr to thnt (mployed in preyious yenls The beetles were confined in hla cardboard boxes of approximately 1 cubic foot capacity Illld to absorb excess moisture the bottoms w(1e lined with fresh moss Apples were used to f(ed the beetles and were l(phnisbed on alternate dllYS After 12 dnys allliye bC(tles which were (lvid(ntly Ullpn1llsitized W(Ie destroy(d nnd the dead olles placed in opm trnys to allow the Atlapsilia which W(lre still ill the Inrvn1 stngr to pupntt Artel pupntion til( UllCOIlshySUlllNl body rontrnts of th( host wrre rellloved to diminnt( decomshyposition gases that might arise durillg shipmrnt to til( detriment of the pllparia The pupnrin v(1e packed betwNm ltl~nS of fresh sphagnum moss in smull vrlltillLted 1ectnnglllnJ tillS and n number of tlllse en tirely surroUlHlpd by 1 3-illClL layer of fllsh mo (1(
pfldwl in illS(Ctploof woodell box(s for shipment One shipnwnt eonsisting of 3ROO pupnrin was SPIlt to tli( rnited
Stnt(s in 1929 This arrind at its dCstinntion Xonmb(r 9 Illvinp bN1l ell rou tp 42 ditys nt it t(mp(rntue of 40deg to 50 0 F
PARASITES OF TilE LAIVA
nEXIlDAE
DEXIA EXTnTIS Aldrich
Delin 1flltrali tilr most rommon of the ChmltHs( dlxiids Wiltshyfirst found Ilt SUiglll ChosllI in 1922 (9) Owing to its (olllplirntld biology no r(Hring work for the shipnwnt of pnmsitiz(d glU bs to t1H lnit(gtd StatN was aWmpted until 1)21) (10) In that Yenr P()pilli(t jnponic(L grubs whirh do not orcur in Chosen (1( eol1letrd filld shipped from nortlH11l Japan to Fukuoka in Kyusitu soutitNll Jnpitn for pnrnsitizlltion by Duia (mnks fonnumiddottkd from Chos) By thr tinlP n suiJici(nt numb(middot of glflYid f(BmTes Wtl( nvailablr Inte i11 JU11r how(yrr most of the grubs w(re too far d(vdop(d for pnrnshysitiztltion purPOR(S find only 850 pnrnsitizld grubs were shipped to til( enitrd States
In 1926 it WfiS d((idpd to (nlTY on rlaring work in 0hos1 utilizing the nutiY( ChOSlll(RC host grubs This provrd to be very sucressful and OWl 20000 grubs were pamsitizrd flnd shippld to titC rnit((l States in 102(3 lind 1927 B(ctlUSC oJ til( dlLngPl iIlYoiwcl in shipping to the rnited Statts Jive grubs othrl thtlll tilos( of POJlillia iapflnica owing to tIl( risk of 111 rscnpl of thrse sp(cils no l(uring w(lIk WIlf
cHrrild on in Chosen in 1928 but 11 mdhod WIIS dlvCloppd whplpby adult fCmale DIJia could be sitipPllaquo LO Yokollilma SII(clssfully and in 1029 rrnring work wns rnrri(d on til(lC the grll bs of P japof ira being IIs((I tiS hosts
Tabl( 2 shows the slllTinl of ndult f(malCs of [Jain middot(lltr(1Ii forshywllrd((l from SlIig(Il Chospn to YokohfilJ11L in 1029 Hnl 1930 til( yciUS that rCaring work WilS el1JTi(d OIl TIle I1dults PI( ronfillNI in thl shipping containl1s which WCI( similm to thos llIlployltd ill transporting Scoliidne npprox11llat(ly 7 days from the timr o( (oll(eshytion to the tim( of removaL
IXVESTIGATIOXS OF PARASITES OF TAPANK~ BEETLE 9
TABLE 2-SuTviLal of adult females of DeJia ventralis shipped from 81tigcn Chosen 10 Yokohama Japa11in 1929 and 1930
Adults Adults surllingYear Generntion shipped shipment
Numb lrumber PercentI
1929_ bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull _ FirsLbullbullbullbullbullbullbullbullbullbullbullbull _bull __ bull __ bullbull_1 063 552 833 1921 __ bullbullbullbullbull ______ bull __ bullbull ____ lhird____ __ 2281 1800 831 1930 __ _bullbullbullbullbullbull __ bull __ First _______ __ ~9() 6S- 68 1910_ _ __ _____ bull_____ __ 1 erond_____ __ __ 2051 14il 720 1930bullbullbull __ bullbullbullbull __ Third __ 2483 2008 809
Total __ __ __ ___ 8368 t 661 ]----()-1 I
It was extremrly difficult to obtain Popillia grubs in any numbers in the --icinity of Yokohamn for rearing work with Deria but this was OYerC0111( by collecting ancl shipping grubs as IHeclNI from Hokshykaido in nortlltrn Japall to Yokohallla n procedure which proyrd to be yery satisfactory and inrxplnsiYe ~0 rearing work was attempted with the s(cond gel1lrntion of DeJia in 1929 oing to the impossibility of s(curing u sufficient number of grubs in IIokkuido during August This was the YNlr of grub scarcity in northern Japall vhen less than 1 percent of the oYCrwintering grubs huying a 2-y(middotnr lifc cyclc (le still in the brynl stage at thn t time of the yeal
The ptlrasitizNl grubs obtain(d tluough parasitization with the first and s(concl g(l1Plntions of DU(L in June nnd August lPspCctiYely wele shipped to thpCnitltC1 Stat(s immpdiatrly aftC1 btinf pnra~itizcd DcYClopment of the parasites took pla(e P11 route nnd the adults emrTgecl sho1tl)- uiter nrriyal at tlHlT destination Grubs containing tbe hibernnting ln1yue of De ia obtuined through pnrasitizatiOIl with the third-g(lHlntion 1Ii(s in 8(ptp111her nnd Octol)(r Pre kPpt OY(1 wintpr at 10kohi1111tl in n wlll-crain(c plot (oY(INI with sod The winttr lllortnlity of these grubs was rnthel high ns only 171 and HL9 pcrcent werc nIl( when dug up [or shipnlPnt in the springs of 1930 lllHl 1931 rlspertiwly
Rwring and Shipping Jfpoundt7wds
The rcuring work with lJJ(l l(fltl(llj1 was accomplisll(d hy using a bnmiddotpding-jnl 111Cthod simillll to that employrd in ]Jle-iolls y(nrs Approximately 50 grubs (Ie placfll in the lonrhalf of a l-qualmiddott glnss jar with soil A fin(-IJ1Psh SCIepn cngp contllining 5 or () lJ(tia f(malp WIlS fittNI into this jur lind rPRtpd on til( soil nnd through this scnPI1 into the soil til( dlpositcd lalTHP pnsNI in palch of the host gruhs 1llP grubs wpl(~ (xtunilHd pneh chty Hlld thORP showing thf pnrnsit( nmiddotpjlntol~ funlwl (1( ((monel nne Tplacpd with frlsh g-rub The flips which Pl( fpd IPg III ll[l with it solution of sunr und watN linmiddotd f[om 3 middot(middotCk tc) J IJ1fll1th in til( cage nnd pnrnitizld flom a5 to )0 pP[cpnt of tlH grub Iach dnv
In pucking the pnrnsitizplt1 gruh 101 shipnwnt iin-lillpd hoxls 1 foot squnrp und Ii inch(s dNp W(IP wwd Each l)()x contnilpd 5 cro-5lction s(ts RPparutd b lin pintp ploviding it COll1pnltllHnt of nhout 1 (lbic inch foJ Pilch of jOIl glubs Fol shipllllllt 2 of thpse hoxps (1( packlrl togtlwl in n (Tong woodpl1 (nsp TIl( soil in which thp gru bs (n pu(k(d had hren thorough Iy fumign tld wi t h (arbon (hsulfid( to (limirmtC any otlw1 inlcl lllntrrial which might htwe
23-13-1----40-2
]0 TECH~ICAL BULLETIN 738 U S DEPT OF AGRICULTURE
been present In table 3 is a summary of shipments of palasitized grubs and the percentages of parasite emcrgenee after arrivul in thl United Stntls
TABLE 3-SlImmary oj shipments oj Popillia japonica grubs l)ara~itizcd willi Deda Ientralis Jrom Japan to the United States 1929-31
Year Contents oC shipment Grubs Parnsites emergingship(led-------1--------------- ---- ______ lVumber lVlLmber I Pentnt
19211-bullbullbullbullbullbullbullbullbullbull_bullbull Grubs pnmsiti1ed with flrstgenerntion Dezia ~ ~500 ~51 1000 1930_bullbullbull __ bull __ bullbullbullbullbull __ Overwintered grubs parnsitized with thirdmiddot 3--
gcnemtionDaiu ~~_~ __ ~~ ___ _ ___ ~ 3000 II 126 1930bullbullbull ___bullbullbullbullbull __bull ___ Grubs pamsitized with lirst-enerntion Detia~~ 8000 1504 18S 1930_____________ ~ _ __ Orubs pllrasitized with ~econd-generntion Drtia 7000 l42J I 203 1930____ __ ~ ~ ~ _ GrUbs pllmsiti1ed with second-genemtion 2500 399 16 (f
Dufu II1931___________ Oerwintered grubs paraSitized with third- 4000 345 8 ( generation Duia I
I 1--------- shyTotaJorneruge~_i- ___ bullbullbullbull __ _ _______ bull ___ ~ ~ ~)(JOOO I 4499middot 155 f i i
Owing to the establishment of Diria lentrali~ in tIl( United Stutes from which materinl for lccolonizlltion was availabll no shipments of this pfllflsitl Wlre made aftlr 1931
PnOSENA SIBII1TA (Fabricins)
Thl dexiid fly Prosena sibi7ita middotliich IS glllclfllly distrihut(d throughout till Asiatic and ~IalaYlln Hegions is an importnnt pilll1shy
sill of PopWia iaponica in Kolth(1l1 Japan It hns olle glllelation a YfIU ami is tllpubl( of palllsitizing and dfwloping Oll numplou sppcils of sealt1bnpid grubs of suitnblr size The life history nnd ililbits of this dexiicl huve been fullv dPidt with in previolls publieashytions (9 10) bull
The shipments from 1923 to H)27 inclusi-e cOl1[-iistN[ of fitldshycolkrtfd grubs of Popillia iap(mit(l whieh showld a getHrnI fitld parflsitization of 10 to 15 percpnt Becaust of till impossibility of detCrting the plesenee of the pUlllsite latnt whieh is nttuchNI to tlH main trnelwal trunk of the host all collpcted glUbs W(I( shipp((L Tllf grubs (re dug in Hoklmido rudy in the spring lind imnHdilltely forwarded to the rnited States arriving thete shortly beforp tll( emlIgenee of the parasites Although this method proed sntisshyfactory the trllnsportntioll costs wPle (xtrCll1(ly high nnd in 1927 it was dfcided to palllsitize till grubs nrtifjeiall)- by 11 mltthod that hnd bCll1 laquo(eloped in 1921 This ms aecomplished by dissecting til( first-stag( larval from tIle oVlsacs of the glUyid femuks and trilllSshy
ferring them to thC host grubs by mcans of a fine blllsh To instlll penetration by the parnsites the grubs with til( smal lnlYllc wIe vlaeed in cclIs of suitable size in wooden bloek fotmiddot it petiot of 24 hours brron being plaecd in thc field whftP tllr oYClwintrlld
Late in the summer of 1928 40596 grubs W(Il artificially PUIllshysitized in 1l0rthCrTJ Japan llnd placed in a prcpnred soil plot w(11 drained coered with sod and surrounded to a depth of 18 inch(s by a fine-mesh screen to pr(vent the gnlbs from wumlcring nwny T11(s( grubs were dug up carly in May 1929 and 19440 were nliv( of whieh 19000 weIpound shipped to the U ni ted Stat(s
In 1929 it was impossibl( to seure sufficient grubs for inoeulatioll work Consequently it was necessary to colleet Prosena femulCs and
IXETrGATIOXS OF PARASITES 01 rAlANFJSFJ DEETLB 11
ship them to the Yokohnma laboTfitory und carryon palasitization work there with available current-year grubs from that vicinity Only 12 p(rcent of the females were alive on arrival at Yokohama Tlus high mortality WitS due to the extremely hot weather prevailing at the time of shipment Out of 364 females received alive 159 contained Inrvae sufficiently developed for this work During the latter part of August 6300 grubs were parasitized and placed in hibernation qunrters As e~perienced with the overwinter~d grubs parasitized with Dexia 1entralis the wintPI mortality was (xceedingly high in this material and only 900 were alive wh(n dug up the followshying spring These with the 19000 shipped in 1929 together rith the 29000 of DeJia tentralis make a total of 48900 grubs containing dexiids thnt were shipped to the United States during the period covcl(d by this bulletin The method of pl1cking tIl( pnrasitized grubs for shipment was the sam( I1S that described for shipment of grubs parnsitized with D ~entrais
No shipments of Prosena sibirita wer( mude aftN 1930 ns this parasite was then established in the Vnit((1 States
SCOLIIDAE
During the course of the investigations from 1920 to 1933 inclusive on the pnrasites of Popillia and related scarabaeid b(etl(s in the Orient a huge number of Scoliidne principally of the genus Tiphia were found and intensively studied ~fost of thtsP species which were consid(recl to be ntw have since been des(ribtd (2 18 19 2021)
In table 4 is presented the 1llown distribution in Tnpllll ChOS(D China Indin nncI Taiwan of all species of Tiphia real(d or shipped sinet the initilltion of the work Of the 29 sp(ei(s list(ci in this tablr 8 nre known to occur ill Japan Chosrn and China 2 in Jupnll nnd ChOS(Il and 1 in ChOS(1l and Chinn No doubt furtlH1 innstigtltions will show a much wider dissemination throughout the Orient Of the species mentioned in this bulletin only liphia bisfculata nnd T tegitiplaga purnsites of Anomala allld occurring in southern Jnpnn have 2 distinct generations a year
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
j
OQ 13
0 0 lts ~
lt ~ a ~ ~ ~ ~
gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
8 TECHXICAL BULLETIN 738 U S DEPT OF GRICFLTCRE
1929 th( majority being brought in from August 1 to 9 th( p(liod when the synchronization of host and parasite is at its best DlIling this time 553885 beetles we1( brought in by the llum(IOtls l1atic collectors FIOIU these 4310 Adapsilia pupalin were obtained giving a parasitization of 078 percent as compared with 053 in 1928
TIlt manuel of renring uncI shipping wus similnr to thnt (mployed in preyious yenls The beetles were confined in hla cardboard boxes of approximately 1 cubic foot capacity Illld to absorb excess moisture the bottoms w(1e lined with fresh moss Apples were used to f(ed the beetles and were l(phnisbed on alternate dllYS After 12 dnys allliye bC(tles which were (lvid(ntly Ullpn1llsitized W(Ie destroy(d nnd the dead olles placed in opm trnys to allow the Atlapsilia which W(lre still ill the Inrvn1 stngr to pupntt Artel pupntion til( UllCOIlshySUlllNl body rontrnts of th( host wrre rellloved to diminnt( decomshyposition gases that might arise durillg shipmrnt to til( detriment of the pllparia The pupnrin v(1e packed betwNm ltl~nS of fresh sphagnum moss in smull vrlltillLted 1ectnnglllnJ tillS and n number of tlllse en tirely surroUlHlpd by 1 3-illClL layer of fllsh mo (1(
pfldwl in illS(Ctploof woodell box(s for shipment One shipnwnt eonsisting of 3ROO pupnrin was SPIlt to tli( rnited
Stnt(s in 1929 This arrind at its dCstinntion Xonmb(r 9 Illvinp bN1l ell rou tp 42 ditys nt it t(mp(rntue of 40deg to 50 0 F
PARASITES OF TilE LAIVA
nEXIlDAE
DEXIA EXTnTIS Aldrich
Delin 1flltrali tilr most rommon of the ChmltHs( dlxiids Wiltshyfirst found Ilt SUiglll ChosllI in 1922 (9) Owing to its (olllplirntld biology no r(Hring work for the shipnwnt of pnmsitiz(d glU bs to t1H lnit(gtd StatN was aWmpted until 1)21) (10) In that Yenr P()pilli(t jnponic(L grubs whirh do not orcur in Chosen (1( eol1letrd filld shipped from nortlH11l Japan to Fukuoka in Kyusitu soutitNll Jnpitn for pnrnsitizlltion by Duia (mnks fonnumiddottkd from Chos) By thr tinlP n suiJici(nt numb(middot of glflYid f(BmTes Wtl( nvailablr Inte i11 JU11r how(yrr most of the grubs w(re too far d(vdop(d for pnrnshysitiztltion purPOR(S find only 850 pnrnsitizld grubs were shipped to til( enitrd States
In 1926 it WfiS d((idpd to (nlTY on rlaring work in 0hos1 utilizing the nutiY( ChOSlll(RC host grubs This provrd to be very sucressful and OWl 20000 grubs were pamsitizrd flnd shippld to titC rnit((l States in 102(3 lind 1927 B(ctlUSC oJ til( dlLngPl iIlYoiwcl in shipping to the rnited Statts Jive grubs othrl thtlll tilos( of POJlillia iapflnica owing to tIl( risk of 111 rscnpl of thrse sp(cils no l(uring w(lIk WIlf
cHrrild on in Chosen in 1928 but 11 mdhod WIIS dlvCloppd whplpby adult fCmale DIJia could be sitipPllaquo LO Yokollilma SII(clssfully and in 1029 rrnring work wns rnrri(d on til(lC the grll bs of P japof ira being IIs((I tiS hosts
Tabl( 2 shows the slllTinl of ndult f(malCs of [Jain middot(lltr(1Ii forshywllrd((l from SlIig(Il Chospn to YokohfilJ11L in 1029 Hnl 1930 til( yciUS that rCaring work WilS el1JTi(d OIl TIle I1dults PI( ronfillNI in thl shipping containl1s which WCI( similm to thos llIlployltd ill transporting Scoliidne npprox11llat(ly 7 days from the timr o( (oll(eshytion to the tim( of removaL
IXVESTIGATIOXS OF PARASITES OF TAPANK~ BEETLE 9
TABLE 2-SuTviLal of adult females of DeJia ventralis shipped from 81tigcn Chosen 10 Yokohama Japa11in 1929 and 1930
Adults Adults surllingYear Generntion shipped shipment
Numb lrumber PercentI
1929_ bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull _ FirsLbullbullbullbullbullbullbullbullbullbullbullbull _bull __ bull __ bullbull_1 063 552 833 1921 __ bullbullbullbullbull ______ bull __ bullbull ____ lhird____ __ 2281 1800 831 1930 __ _bullbullbullbullbullbull __ bull __ First _______ __ ~9() 6S- 68 1910_ _ __ _____ bull_____ __ 1 erond_____ __ __ 2051 14il 720 1930bullbullbull __ bullbullbullbull __ Third __ 2483 2008 809
Total __ __ __ ___ 8368 t 661 ]----()-1 I
It was extremrly difficult to obtain Popillia grubs in any numbers in the --icinity of Yokohamn for rearing work with Deria but this was OYerC0111( by collecting ancl shipping grubs as IHeclNI from Hokshykaido in nortlltrn Japall to Yokohallla n procedure which proyrd to be yery satisfactory and inrxplnsiYe ~0 rearing work was attempted with the s(cond gel1lrntion of DeJia in 1929 oing to the impossibility of s(curing u sufficient number of grubs in IIokkuido during August This was the YNlr of grub scarcity in northern Japall vhen less than 1 percent of the oYCrwintering grubs huying a 2-y(middotnr lifc cyclc (le still in the brynl stage at thn t time of the yeal
The ptlrasitizNl grubs obtain(d tluough parasitization with the first and s(concl g(l1Plntions of DU(L in June nnd August lPspCctiYely wele shipped to thpCnitltC1 Stat(s immpdiatrly aftC1 btinf pnra~itizcd DcYClopment of the parasites took pla(e P11 route nnd the adults emrTgecl sho1tl)- uiter nrriyal at tlHlT destination Grubs containing tbe hibernnting ln1yue of De ia obtuined through pnrasitizatiOIl with the third-g(lHlntion 1Ii(s in 8(ptp111her nnd Octol)(r Pre kPpt OY(1 wintpr at 10kohi1111tl in n wlll-crain(c plot (oY(INI with sod The winttr lllortnlity of these grubs was rnthel high ns only 171 and HL9 pcrcent werc nIl( when dug up [or shipnlPnt in the springs of 1930 lllHl 1931 rlspertiwly
Rwring and Shipping Jfpoundt7wds
The rcuring work with lJJ(l l(fltl(llj1 was accomplisll(d hy using a bnmiddotpding-jnl 111Cthod simillll to that employrd in ]Jle-iolls y(nrs Approximately 50 grubs (Ie placfll in the lonrhalf of a l-qualmiddott glnss jar with soil A fin(-IJ1Psh SCIepn cngp contllining 5 or () lJ(tia f(malp WIlS fittNI into this jur lind rPRtpd on til( soil nnd through this scnPI1 into the soil til( dlpositcd lalTHP pnsNI in palch of the host gruhs 1llP grubs wpl(~ (xtunilHd pneh chty Hlld thORP showing thf pnrnsit( nmiddotpjlntol~ funlwl (1( ((monel nne Tplacpd with frlsh g-rub The flips which Pl( fpd IPg III ll[l with it solution of sunr und watN linmiddotd f[om 3 middot(middotCk tc) J IJ1fll1th in til( cage nnd pnrnitizld flom a5 to )0 pP[cpnt of tlH grub Iach dnv
In pucking the pnrnsitizplt1 gruh 101 shipnwnt iin-lillpd hoxls 1 foot squnrp und Ii inch(s dNp W(IP wwd Each l)()x contnilpd 5 cro-5lction s(ts RPparutd b lin pintp ploviding it COll1pnltllHnt of nhout 1 (lbic inch foJ Pilch of jOIl glubs Fol shipllllllt 2 of thpse hoxps (1( packlrl togtlwl in n (Tong woodpl1 (nsp TIl( soil in which thp gru bs (n pu(k(d had hren thorough Iy fumign tld wi t h (arbon (hsulfid( to (limirmtC any otlw1 inlcl lllntrrial which might htwe
23-13-1----40-2
]0 TECH~ICAL BULLETIN 738 U S DEPT OF AGRICULTURE
been present In table 3 is a summary of shipments of palasitized grubs and the percentages of parasite emcrgenee after arrivul in thl United Stntls
TABLE 3-SlImmary oj shipments oj Popillia japonica grubs l)ara~itizcd willi Deda Ientralis Jrom Japan to the United States 1929-31
Year Contents oC shipment Grubs Parnsites emergingship(led-------1--------------- ---- ______ lVumber lVlLmber I Pentnt
19211-bullbullbullbullbullbullbullbullbullbull_bullbull Grubs pnmsiti1ed with flrstgenerntion Dezia ~ ~500 ~51 1000 1930_bullbullbull __ bull __ bullbullbullbullbull __ Overwintered grubs parnsitized with thirdmiddot 3--
gcnemtionDaiu ~~_~ __ ~~ ___ _ ___ ~ 3000 II 126 1930bullbullbull ___bullbullbullbullbull __bull ___ Grubs pamsitized with lirst-enerntion Detia~~ 8000 1504 18S 1930_____________ ~ _ __ Orubs pllrasitized with ~econd-generntion Drtia 7000 l42J I 203 1930____ __ ~ ~ ~ _ GrUbs pllmsiti1ed with second-genemtion 2500 399 16 (f
Dufu II1931___________ Oerwintered grubs paraSitized with third- 4000 345 8 ( generation Duia I
I 1--------- shyTotaJorneruge~_i- ___ bullbullbullbull __ _ _______ bull ___ ~ ~ ~)(JOOO I 4499middot 155 f i i
Owing to the establishment of Diria lentrali~ in tIl( United Stutes from which materinl for lccolonizlltion was availabll no shipments of this pfllflsitl Wlre made aftlr 1931
PnOSENA SIBII1TA (Fabricins)
Thl dexiid fly Prosena sibi7ita middotliich IS glllclfllly distrihut(d throughout till Asiatic and ~IalaYlln Hegions is an importnnt pilll1shy
sill of PopWia iaponica in Kolth(1l1 Japan It hns olle glllelation a YfIU ami is tllpubl( of palllsitizing and dfwloping Oll numplou sppcils of sealt1bnpid grubs of suitnblr size The life history nnd ililbits of this dexiicl huve been fullv dPidt with in previolls publieashytions (9 10) bull
The shipments from 1923 to H)27 inclusi-e cOl1[-iistN[ of fitldshycolkrtfd grubs of Popillia iap(mit(l whieh showld a getHrnI fitld parflsitization of 10 to 15 percpnt Becaust of till impossibility of detCrting the plesenee of the pUlllsite latnt whieh is nttuchNI to tlH main trnelwal trunk of the host all collpcted glUbs W(I( shipp((L Tllf grubs (re dug in Hoklmido rudy in the spring lind imnHdilltely forwarded to the rnited States arriving thete shortly beforp tll( emlIgenee of the parasites Although this method proed sntisshyfactory the trllnsportntioll costs wPle (xtrCll1(ly high nnd in 1927 it was dfcided to palllsitize till grubs nrtifjeiall)- by 11 mltthod that hnd bCll1 laquo(eloped in 1921 This ms aecomplished by dissecting til( first-stag( larval from tIle oVlsacs of the glUyid femuks and trilllSshy
ferring them to thC host grubs by mcans of a fine blllsh To instlll penetration by the parnsites the grubs with til( smal lnlYllc wIe vlaeed in cclIs of suitable size in wooden bloek fotmiddot it petiot of 24 hours brron being plaecd in thc field whftP tllr oYClwintrlld
Late in the summer of 1928 40596 grubs W(Il artificially PUIllshysitized in 1l0rthCrTJ Japan llnd placed in a prcpnred soil plot w(11 drained coered with sod and surrounded to a depth of 18 inch(s by a fine-mesh screen to pr(vent the gnlbs from wumlcring nwny T11(s( grubs were dug up carly in May 1929 and 19440 were nliv( of whieh 19000 weIpound shipped to the U ni ted Stat(s
In 1929 it was impossibl( to seure sufficient grubs for inoeulatioll work Consequently it was necessary to colleet Prosena femulCs and
IXETrGATIOXS OF PARASITES 01 rAlANFJSFJ DEETLB 11
ship them to the Yokohnma laboTfitory und carryon palasitization work there with available current-year grubs from that vicinity Only 12 p(rcent of the females were alive on arrival at Yokohama Tlus high mortality WitS due to the extremely hot weather prevailing at the time of shipment Out of 364 females received alive 159 contained Inrvae sufficiently developed for this work During the latter part of August 6300 grubs were parasitized and placed in hibernation qunrters As e~perienced with the overwinter~d grubs parasitized with Dexia 1entralis the wintPI mortality was (xceedingly high in this material and only 900 were alive wh(n dug up the followshying spring These with the 19000 shipped in 1929 together rith the 29000 of DeJia tentralis make a total of 48900 grubs containing dexiids thnt were shipped to the United States during the period covcl(d by this bulletin The method of pl1cking tIl( pnrasitized grubs for shipment was the sam( I1S that described for shipment of grubs parnsitized with D ~entrais
No shipments of Prosena sibirita wer( mude aftN 1930 ns this parasite was then established in the Vnit((1 States
SCOLIIDAE
During the course of the investigations from 1920 to 1933 inclusive on the pnrasites of Popillia and related scarabaeid b(etl(s in the Orient a huge number of Scoliidne principally of the genus Tiphia were found and intensively studied ~fost of thtsP species which were consid(recl to be ntw have since been des(ribtd (2 18 19 2021)
In table 4 is presented the 1llown distribution in Tnpllll ChOS(D China Indin nncI Taiwan of all species of Tiphia real(d or shipped sinet the initilltion of the work Of the 29 sp(ei(s list(ci in this tablr 8 nre known to occur ill Japan Chosrn and China 2 in Jupnll nnd ChOS(Il and 1 in ChOS(1l and Chinn No doubt furtlH1 innstigtltions will show a much wider dissemination throughout the Orient Of the species mentioned in this bulletin only liphia bisfculata nnd T tegitiplaga purnsites of Anomala allld occurring in southern Jnpnn have 2 distinct generations a year
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
j
OQ 13
0 0 lts ~
lt ~ a ~ ~ ~ ~
gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
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34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
IXVESTIGATIOXS OF PARASITES OF TAPANK~ BEETLE 9
TABLE 2-SuTviLal of adult females of DeJia ventralis shipped from 81tigcn Chosen 10 Yokohama Japa11in 1929 and 1930
Adults Adults surllingYear Generntion shipped shipment
Numb lrumber PercentI
1929_ bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull _ FirsLbullbullbullbullbullbullbullbullbullbullbullbull _bull __ bull __ bullbull_1 063 552 833 1921 __ bullbullbullbullbull ______ bull __ bullbull ____ lhird____ __ 2281 1800 831 1930 __ _bullbullbullbullbullbull __ bull __ First _______ __ ~9() 6S- 68 1910_ _ __ _____ bull_____ __ 1 erond_____ __ __ 2051 14il 720 1930bullbullbull __ bullbullbullbull __ Third __ 2483 2008 809
Total __ __ __ ___ 8368 t 661 ]----()-1 I
It was extremrly difficult to obtain Popillia grubs in any numbers in the --icinity of Yokohamn for rearing work with Deria but this was OYerC0111( by collecting ancl shipping grubs as IHeclNI from Hokshykaido in nortlltrn Japall to Yokohallla n procedure which proyrd to be yery satisfactory and inrxplnsiYe ~0 rearing work was attempted with the s(cond gel1lrntion of DeJia in 1929 oing to the impossibility of s(curing u sufficient number of grubs in IIokkuido during August This was the YNlr of grub scarcity in northern Japall vhen less than 1 percent of the oYCrwintering grubs huying a 2-y(middotnr lifc cyclc (le still in the brynl stage at thn t time of the yeal
The ptlrasitizNl grubs obtain(d tluough parasitization with the first and s(concl g(l1Plntions of DU(L in June nnd August lPspCctiYely wele shipped to thpCnitltC1 Stat(s immpdiatrly aftC1 btinf pnra~itizcd DcYClopment of the parasites took pla(e P11 route nnd the adults emrTgecl sho1tl)- uiter nrriyal at tlHlT destination Grubs containing tbe hibernnting ln1yue of De ia obtuined through pnrasitizatiOIl with the third-g(lHlntion 1Ii(s in 8(ptp111her nnd Octol)(r Pre kPpt OY(1 wintpr at 10kohi1111tl in n wlll-crain(c plot (oY(INI with sod The winttr lllortnlity of these grubs was rnthel high ns only 171 and HL9 pcrcent werc nIl( when dug up [or shipnlPnt in the springs of 1930 lllHl 1931 rlspertiwly
Rwring and Shipping Jfpoundt7wds
The rcuring work with lJJ(l l(fltl(llj1 was accomplisll(d hy using a bnmiddotpding-jnl 111Cthod simillll to that employrd in ]Jle-iolls y(nrs Approximately 50 grubs (Ie placfll in the lonrhalf of a l-qualmiddott glnss jar with soil A fin(-IJ1Psh SCIepn cngp contllining 5 or () lJ(tia f(malp WIlS fittNI into this jur lind rPRtpd on til( soil nnd through this scnPI1 into the soil til( dlpositcd lalTHP pnsNI in palch of the host gruhs 1llP grubs wpl(~ (xtunilHd pneh chty Hlld thORP showing thf pnrnsit( nmiddotpjlntol~ funlwl (1( ((monel nne Tplacpd with frlsh g-rub The flips which Pl( fpd IPg III ll[l with it solution of sunr und watN linmiddotd f[om 3 middot(middotCk tc) J IJ1fll1th in til( cage nnd pnrnitizld flom a5 to )0 pP[cpnt of tlH grub Iach dnv
In pucking the pnrnsitizplt1 gruh 101 shipnwnt iin-lillpd hoxls 1 foot squnrp und Ii inch(s dNp W(IP wwd Each l)()x contnilpd 5 cro-5lction s(ts RPparutd b lin pintp ploviding it COll1pnltllHnt of nhout 1 (lbic inch foJ Pilch of jOIl glubs Fol shipllllllt 2 of thpse hoxps (1( packlrl togtlwl in n (Tong woodpl1 (nsp TIl( soil in which thp gru bs (n pu(k(d had hren thorough Iy fumign tld wi t h (arbon (hsulfid( to (limirmtC any otlw1 inlcl lllntrrial which might htwe
23-13-1----40-2
]0 TECH~ICAL BULLETIN 738 U S DEPT OF AGRICULTURE
been present In table 3 is a summary of shipments of palasitized grubs and the percentages of parasite emcrgenee after arrivul in thl United Stntls
TABLE 3-SlImmary oj shipments oj Popillia japonica grubs l)ara~itizcd willi Deda Ientralis Jrom Japan to the United States 1929-31
Year Contents oC shipment Grubs Parnsites emergingship(led-------1--------------- ---- ______ lVumber lVlLmber I Pentnt
19211-bullbullbullbullbullbullbullbullbullbull_bullbull Grubs pnmsiti1ed with flrstgenerntion Dezia ~ ~500 ~51 1000 1930_bullbullbull __ bull __ bullbullbullbullbull __ Overwintered grubs parnsitized with thirdmiddot 3--
gcnemtionDaiu ~~_~ __ ~~ ___ _ ___ ~ 3000 II 126 1930bullbullbull ___bullbullbullbullbull __bull ___ Grubs pamsitized with lirst-enerntion Detia~~ 8000 1504 18S 1930_____________ ~ _ __ Orubs pllrasitized with ~econd-generntion Drtia 7000 l42J I 203 1930____ __ ~ ~ ~ _ GrUbs pllmsiti1ed with second-genemtion 2500 399 16 (f
Dufu II1931___________ Oerwintered grubs paraSitized with third- 4000 345 8 ( generation Duia I
I 1--------- shyTotaJorneruge~_i- ___ bullbullbullbull __ _ _______ bull ___ ~ ~ ~)(JOOO I 4499middot 155 f i i
Owing to the establishment of Diria lentrali~ in tIl( United Stutes from which materinl for lccolonizlltion was availabll no shipments of this pfllflsitl Wlre made aftlr 1931
PnOSENA SIBII1TA (Fabricins)
Thl dexiid fly Prosena sibi7ita middotliich IS glllclfllly distrihut(d throughout till Asiatic and ~IalaYlln Hegions is an importnnt pilll1shy
sill of PopWia iaponica in Kolth(1l1 Japan It hns olle glllelation a YfIU ami is tllpubl( of palllsitizing and dfwloping Oll numplou sppcils of sealt1bnpid grubs of suitnblr size The life history nnd ililbits of this dexiicl huve been fullv dPidt with in previolls publieashytions (9 10) bull
The shipments from 1923 to H)27 inclusi-e cOl1[-iistN[ of fitldshycolkrtfd grubs of Popillia iap(mit(l whieh showld a getHrnI fitld parflsitization of 10 to 15 percpnt Becaust of till impossibility of detCrting the plesenee of the pUlllsite latnt whieh is nttuchNI to tlH main trnelwal trunk of the host all collpcted glUbs W(I( shipp((L Tllf grubs (re dug in Hoklmido rudy in the spring lind imnHdilltely forwarded to the rnited States arriving thete shortly beforp tll( emlIgenee of the parasites Although this method proed sntisshyfactory the trllnsportntioll costs wPle (xtrCll1(ly high nnd in 1927 it was dfcided to palllsitize till grubs nrtifjeiall)- by 11 mltthod that hnd bCll1 laquo(eloped in 1921 This ms aecomplished by dissecting til( first-stag( larval from tIle oVlsacs of the glUyid femuks and trilllSshy
ferring them to thC host grubs by mcans of a fine blllsh To instlll penetration by the parnsites the grubs with til( smal lnlYllc wIe vlaeed in cclIs of suitable size in wooden bloek fotmiddot it petiot of 24 hours brron being plaecd in thc field whftP tllr oYClwintrlld
Late in the summer of 1928 40596 grubs W(Il artificially PUIllshysitized in 1l0rthCrTJ Japan llnd placed in a prcpnred soil plot w(11 drained coered with sod and surrounded to a depth of 18 inch(s by a fine-mesh screen to pr(vent the gnlbs from wumlcring nwny T11(s( grubs were dug up carly in May 1929 and 19440 were nliv( of whieh 19000 weIpound shipped to the U ni ted Stat(s
In 1929 it was impossibl( to seure sufficient grubs for inoeulatioll work Consequently it was necessary to colleet Prosena femulCs and
IXETrGATIOXS OF PARASITES 01 rAlANFJSFJ DEETLB 11
ship them to the Yokohnma laboTfitory und carryon palasitization work there with available current-year grubs from that vicinity Only 12 p(rcent of the females were alive on arrival at Yokohama Tlus high mortality WitS due to the extremely hot weather prevailing at the time of shipment Out of 364 females received alive 159 contained Inrvae sufficiently developed for this work During the latter part of August 6300 grubs were parasitized and placed in hibernation qunrters As e~perienced with the overwinter~d grubs parasitized with Dexia 1entralis the wintPI mortality was (xceedingly high in this material and only 900 were alive wh(n dug up the followshying spring These with the 19000 shipped in 1929 together rith the 29000 of DeJia tentralis make a total of 48900 grubs containing dexiids thnt were shipped to the United States during the period covcl(d by this bulletin The method of pl1cking tIl( pnrasitized grubs for shipment was the sam( I1S that described for shipment of grubs parnsitized with D ~entrais
No shipments of Prosena sibirita wer( mude aftN 1930 ns this parasite was then established in the Vnit((1 States
SCOLIIDAE
During the course of the investigations from 1920 to 1933 inclusive on the pnrasites of Popillia and related scarabaeid b(etl(s in the Orient a huge number of Scoliidne principally of the genus Tiphia were found and intensively studied ~fost of thtsP species which were consid(recl to be ntw have since been des(ribtd (2 18 19 2021)
In table 4 is presented the 1llown distribution in Tnpllll ChOS(D China Indin nncI Taiwan of all species of Tiphia real(d or shipped sinet the initilltion of the work Of the 29 sp(ei(s list(ci in this tablr 8 nre known to occur ill Japan Chosrn and China 2 in Jupnll nnd ChOS(Il and 1 in ChOS(1l and Chinn No doubt furtlH1 innstigtltions will show a much wider dissemination throughout the Orient Of the species mentioned in this bulletin only liphia bisfculata nnd T tegitiplaga purnsites of Anomala allld occurring in southern Jnpnn have 2 distinct generations a year
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
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gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
]0 TECH~ICAL BULLETIN 738 U S DEPT OF AGRICULTURE
been present In table 3 is a summary of shipments of palasitized grubs and the percentages of parasite emcrgenee after arrivul in thl United Stntls
TABLE 3-SlImmary oj shipments oj Popillia japonica grubs l)ara~itizcd willi Deda Ientralis Jrom Japan to the United States 1929-31
Year Contents oC shipment Grubs Parnsites emergingship(led-------1--------------- ---- ______ lVumber lVlLmber I Pentnt
19211-bullbullbullbullbullbullbullbullbullbull_bullbull Grubs pnmsiti1ed with flrstgenerntion Dezia ~ ~500 ~51 1000 1930_bullbullbull __ bull __ bullbullbullbullbull __ Overwintered grubs parnsitized with thirdmiddot 3--
gcnemtionDaiu ~~_~ __ ~~ ___ _ ___ ~ 3000 II 126 1930bullbullbull ___bullbullbullbullbull __bull ___ Grubs pamsitized with lirst-enerntion Detia~~ 8000 1504 18S 1930_____________ ~ _ __ Orubs pllrasitized with ~econd-generntion Drtia 7000 l42J I 203 1930____ __ ~ ~ ~ _ GrUbs pllmsiti1ed with second-genemtion 2500 399 16 (f
Dufu II1931___________ Oerwintered grubs paraSitized with third- 4000 345 8 ( generation Duia I
I 1--------- shyTotaJorneruge~_i- ___ bullbullbullbull __ _ _______ bull ___ ~ ~ ~)(JOOO I 4499middot 155 f i i
Owing to the establishment of Diria lentrali~ in tIl( United Stutes from which materinl for lccolonizlltion was availabll no shipments of this pfllflsitl Wlre made aftlr 1931
PnOSENA SIBII1TA (Fabricins)
Thl dexiid fly Prosena sibi7ita middotliich IS glllclfllly distrihut(d throughout till Asiatic and ~IalaYlln Hegions is an importnnt pilll1shy
sill of PopWia iaponica in Kolth(1l1 Japan It hns olle glllelation a YfIU ami is tllpubl( of palllsitizing and dfwloping Oll numplou sppcils of sealt1bnpid grubs of suitnblr size The life history nnd ililbits of this dexiicl huve been fullv dPidt with in previolls publieashytions (9 10) bull
The shipments from 1923 to H)27 inclusi-e cOl1[-iistN[ of fitldshycolkrtfd grubs of Popillia iap(mit(l whieh showld a getHrnI fitld parflsitization of 10 to 15 percpnt Becaust of till impossibility of detCrting the plesenee of the pUlllsite latnt whieh is nttuchNI to tlH main trnelwal trunk of the host all collpcted glUbs W(I( shipp((L Tllf grubs (re dug in Hoklmido rudy in the spring lind imnHdilltely forwarded to the rnited States arriving thete shortly beforp tll( emlIgenee of the parasites Although this method proed sntisshyfactory the trllnsportntioll costs wPle (xtrCll1(ly high nnd in 1927 it was dfcided to palllsitize till grubs nrtifjeiall)- by 11 mltthod that hnd bCll1 laquo(eloped in 1921 This ms aecomplished by dissecting til( first-stag( larval from tIle oVlsacs of the glUyid femuks and trilllSshy
ferring them to thC host grubs by mcans of a fine blllsh To instlll penetration by the parnsites the grubs with til( smal lnlYllc wIe vlaeed in cclIs of suitable size in wooden bloek fotmiddot it petiot of 24 hours brron being plaecd in thc field whftP tllr oYClwintrlld
Late in the summer of 1928 40596 grubs W(Il artificially PUIllshysitized in 1l0rthCrTJ Japan llnd placed in a prcpnred soil plot w(11 drained coered with sod and surrounded to a depth of 18 inch(s by a fine-mesh screen to pr(vent the gnlbs from wumlcring nwny T11(s( grubs were dug up carly in May 1929 and 19440 were nliv( of whieh 19000 weIpound shipped to the U ni ted Stat(s
In 1929 it was impossibl( to seure sufficient grubs for inoeulatioll work Consequently it was necessary to colleet Prosena femulCs and
IXETrGATIOXS OF PARASITES 01 rAlANFJSFJ DEETLB 11
ship them to the Yokohnma laboTfitory und carryon palasitization work there with available current-year grubs from that vicinity Only 12 p(rcent of the females were alive on arrival at Yokohama Tlus high mortality WitS due to the extremely hot weather prevailing at the time of shipment Out of 364 females received alive 159 contained Inrvae sufficiently developed for this work During the latter part of August 6300 grubs were parasitized and placed in hibernation qunrters As e~perienced with the overwinter~d grubs parasitized with Dexia 1entralis the wintPI mortality was (xceedingly high in this material and only 900 were alive wh(n dug up the followshying spring These with the 19000 shipped in 1929 together rith the 29000 of DeJia tentralis make a total of 48900 grubs containing dexiids thnt were shipped to the United States during the period covcl(d by this bulletin The method of pl1cking tIl( pnrasitized grubs for shipment was the sam( I1S that described for shipment of grubs parnsitized with D ~entrais
No shipments of Prosena sibirita wer( mude aftN 1930 ns this parasite was then established in the Vnit((1 States
SCOLIIDAE
During the course of the investigations from 1920 to 1933 inclusive on the pnrasites of Popillia and related scarabaeid b(etl(s in the Orient a huge number of Scoliidne principally of the genus Tiphia were found and intensively studied ~fost of thtsP species which were consid(recl to be ntw have since been des(ribtd (2 18 19 2021)
In table 4 is presented the 1llown distribution in Tnpllll ChOS(D China Indin nncI Taiwan of all species of Tiphia real(d or shipped sinet the initilltion of the work Of the 29 sp(ei(s list(ci in this tablr 8 nre known to occur ill Japan Chosrn and China 2 in Jupnll nnd ChOS(Il and 1 in ChOS(1l and Chinn No doubt furtlH1 innstigtltions will show a much wider dissemination throughout the Orient Of the species mentioned in this bulletin only liphia bisfculata nnd T tegitiplaga purnsites of Anomala allld occurring in southern Jnpnn have 2 distinct generations a year
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
j
OQ 13
0 0 lts ~
lt ~ a ~ ~ ~ ~
gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
IXETrGATIOXS OF PARASITES 01 rAlANFJSFJ DEETLB 11
ship them to the Yokohnma laboTfitory und carryon palasitization work there with available current-year grubs from that vicinity Only 12 p(rcent of the females were alive on arrival at Yokohama Tlus high mortality WitS due to the extremely hot weather prevailing at the time of shipment Out of 364 females received alive 159 contained Inrvae sufficiently developed for this work During the latter part of August 6300 grubs were parasitized and placed in hibernation qunrters As e~perienced with the overwinter~d grubs parasitized with Dexia 1entralis the wintPI mortality was (xceedingly high in this material and only 900 were alive wh(n dug up the followshying spring These with the 19000 shipped in 1929 together rith the 29000 of DeJia tentralis make a total of 48900 grubs containing dexiids thnt were shipped to the United States during the period covcl(d by this bulletin The method of pl1cking tIl( pnrasitized grubs for shipment was the sam( I1S that described for shipment of grubs parnsitized with D ~entrais
No shipments of Prosena sibirita wer( mude aftN 1930 ns this parasite was then established in the Vnit((1 States
SCOLIIDAE
During the course of the investigations from 1920 to 1933 inclusive on the pnrasites of Popillia and related scarabaeid b(etl(s in the Orient a huge number of Scoliidne principally of the genus Tiphia were found and intensively studied ~fost of thtsP species which were consid(recl to be ntw have since been des(ribtd (2 18 19 2021)
In table 4 is presented the 1llown distribution in Tnpllll ChOS(D China Indin nncI Taiwan of all species of Tiphia real(d or shipped sinet the initilltion of the work Of the 29 sp(ei(s list(ci in this tablr 8 nre known to occur ill Japan Chosrn and China 2 in Jupnll nnd ChOS(Il and 1 in ChOS(1l and Chinn No doubt furtlH1 innstigtltions will show a much wider dissemination throughout the Orient Of the species mentioned in this bulletin only liphia bisfculata nnd T tegitiplaga purnsites of Anomala allld occurring in southern Jnpnn have 2 distinct generations a year
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
j
OQ 13
0 0 lts ~
lt ~ a ~ ~ ~ ~
gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
lAHlE 4-11ecord of knuwn distribution uf various spcies of Piphia in the Orient l)
--- ---- -~~-------~--------~----~
j
OQ 13
0 0 lts ~
lt ~ a ~ ~ ~ ~
gt0 1 c Counlrr tlllltl Ilreftlt(ure I ~ 1 ~ oS ~ 1 e 1 en E-lt E-lt ~ 1t11Jitr t ()r pronnCH t 5 5
gt1 ~ ( i5 2 1 2 amp g go ci sect ~ shy1 ~ g sect ~ ~ c oS ~ ~ 0 ~ 2 2 ~ ~ i
t sect~ ~ ~ ~ Q ~ 1 t E lt ~
1 i5 ~ - ~ ~ ~ ~ s s ~ ~ ~ ~ ~ ~I ~ ~ ~ ~ ~ 1 1 _~ I __ __ __ __ __ __I __ __ __I_I -1- -1----1__1 middot__1--____ ______ _ _______ __ __ ____ __1__ 1__ ~
)11- ___ _ ~_ ___ __ ___ ____ ____ X ____ ____ ____ ____ ___ _lshlkulimiddotkn Sllllpnro_ _~ ~~ ~ Akitu-k11 _ WUtlll _ _ _ xL _____ bull__ _____ _ X ____ _ - bull ____ ---- ____ ___ --- ---- ---- ---- --- shy
______ ___ bull_ ____________ X X __ bull__ bull ____ __ --_ - ___ ---- ---- -- ~ KOiWlli X X XhVHtemiddotklIl _ ~
1liyugl-kcn t traquoII~~~T- -- --_~[ ~ t- -_ -~~ -~j~ -~l ~ ~ --j ~- -j- ~j l
~lJ~allO~~ X ~ - X middotX- iC-1 i Ntl~nl1u-ktIL _
lktlIl1 __ X _ -- -00 bullbullbull - --- shy iflIWlllllllrudll _ _ _ X _ ____ __ X -__ X _ --- --- shyJoyoslI X X X __ _ __ X ---- --- ---bullbull--shy ~ ~~~~~~~ -- ~ ~
loeh igi-kcll ~ ~N~~gIlt X _ -- = ~ = ~ ~ ~ Sllltnlllll-kln o
gtj
lokyo-(u _ tU~iIt~~middotu X -~~ -~~ ~~~~ ~~~~ ~~ ~~ ~~~~ ~~~~ i ~~~~ ~~~ ~~~~ E HhilllllarUkO X bull ---- bullbullbullbullbull--- ---- -- ---shy
III IIsllillloliJ I X
X 1 I X ---- ---- ---- ----~
Knllll~ltn~klU ~ I Yokohlllllll X X I X X X X X X X ---- --- shy11l511shl _ X -- -- (hi~u(lki X ~ I Kllllllllllllliyu I I 1 I I bull X C1
I Nurnshillll I - I Xi I I - X - lhlhlmiddotk1l KlytlWIIII i I I 1 I --- X ~ 1lIho + I 1 X X ___ X __ I - X X - -- x X ------ bull -- shy8h iZllnka-ktll llnmnntntslI bullbull _ _1 JJ1 1 _ I J _11 __ --_ ___ _______-JJ X _ bullbullbull------
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
Y~IIJ1~nushi(lI bull ~Orubullbull middotmiddotmiddotmiddotmiddotmiddotmiddotmiddot1middotmiddotmiddotmiddot 1 x Il--I x -x---=~~~~~~~~~_-- M~~I~ril~middot ~ ~ - ~ ~ I~ ~ ~ = Ntlra-k()J) KoriYUJIlIl ~~~ X ~ -- ~~~ ~ xr-middotmiddotmiddotmiddot J1ivo~kNI H~kk(SII X ~ H
bull Uisuzakl bull X ~ middotllklluknmiddotk(middotJI I(~UhUSh i
jKIlJVtlbullbullbull X I X X ~ III-x-III= 3X (fJOitlklII l ASlji Xshy
lf3eppu bullbullbullbullbull 11x-I 1 1 1-3X Inkigllchi bull bull X H
_1 x xC xKUIlUUtlOto-kOIl ~_~~NlJllino_ shy
X --- x-IIII sectMIImiddotnjl X X ]iYllznki Kohayashi _ ~ -- x X 1middotmiddotmiddotmiddot1middotmiddot middotmiddot-middot X - -bullbull-X __ __ bullbullbull __ X ___ -bull ojKiriShimu X KllgoshimlmiddotkcII r~hik x 2
oshlllllltsu X en ZIn X --11- -1-middotmiddotmiddotmiddotmiddotmiddotI = Nilgasaklmiddotkllli oShhnniJllru X bullbull bull_ lj
(1I0Sgt- Kukuku Shllriill X
H g X X x x xmiddot YIlxlxIY ~ ~ IIX middotxmiddot -X = ~ I(middotiklmiddotlo OI~JI~~-oj~O - __ _ X _ X _ ld __ _ X bullbullbull_ bullbullbull X _ bull _bullbullbull YUjyO xL
enChuscimiddotlIUIlmiddotcO ~(lknn HPnidtJl ~ _ LC X- ~
K(ishllmiddothokumiddot(o _ X _bullbull t3 Z(lnru-rlanmiddotlo Koshu x _ ~ - _ I ~ en
Klmiddotishu
Kclshol1J1middotdo 1IIr1I xlI1==IJ(1II-IlSlIlI (111 o
Chillltlnn~ Killugsu bull lenllill _ _~ +_ ~ x ~ ~ ~ ~ I~ = =
lj
yullgchowbullbullbullbull Szrchusrll
~~iiiclioiv~ lx- -middotmiddot IIIlXmiddot y xmiddot Se y ~ Chekiang ~1Il~po_~ __ _ X _bull _ X bullbull _ bull - - ---Kulillng bullbull middotxmiddot XI X _ _ bull _ bull __ X X _ _ X _ 2l
tjHUnklnng - - - bull _ _ _ _ ___ _ bull __ X __ - enAnwhrl NlIlIklnl1 bull __
~ ~
X _ bull_ bull __ bull X bullbull _ _ - __ - tj _I ~_~I_ ~ _~ ~ ~ ~ bull _ ~_ ~ ~_~~ bullbull ~ _ __ ~ ~w ~_yen_ __ ~ ___ ~ _ ~~ _ _~ __ X ~_~~ ~___ w_~_Ylhzlil _ I -- X _bullbull___ bull _bullbull____ bull X __ __ __ bull_ X _bull _X ~ ltI~ __I middot A ~lIiJ101lNtH SSlllil - Chrrrnflunji _ __ 1middot- 1- ~
tjJIgt lulhoku-lhu JIokulo_ J _middotr- x 11 r Y y
~
~
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
14 TECHNICAL BLLLETIN 738 U S DEPT OF AGRICrLTlmiddotUE
Investigations have shown that the normal host grubs of these species of Tiphia occurring in several countries fall within generic rather than specific limits This may be exemplified by T 1ernalis which has Popillia quaciliguftata as its normal host in Chosen and P (hinensis in China but will oviposit and develop on P Japonica equnlly as weil There is little variation in the biology of those species occurring in different countries or in widely separated areus in the same country other than the time of ftppearance of adults in the field TIllS variution in field appenrance is due to temperature conditions and the presence of suitable grubs in the soil
The luck of sufficient adult food and of suitlble host glubs in the field are limiting factors that influence the effectiveness und distribushytion of TipMa The food supply of the adult consists mainly of honeydew exuded by aphids and other bugs the blossoms of various plants principally Umbelliferae and Polygonaceae and the nectar from glands nssociated with certain plants such as the sweetpotato Suitable host grubs are those species wlllch are of sufficient size and are accepted by the Tiphia and are present in the field usunlly near the parnsite feeding grounds at the time when Tiphia adults appear in the field
The position of the egg on the host grub for each spec if qf Tiphia discussed in this bulletin is illustrated in figure 1 In the previous publications (9 10) the egg position on the host grub is figured for all the species investigated during the period 1920-28 some of which are not mentioned ill this bulletin
Teglplogo 8iseculata Burrelli Sternata Vrnalis
Isolota Pullvora
Hamancularis SOfo
Coston easvora and sp
Popilliovoo
Notopolita aIem
Asericae
Maturo and sp
1ltG(RE I-The position of the eggs of llrious species of Tiphia on the host ~rnb
TIPHlA VERNALIS Rohwer
Tiphia lernalis WI1S first found in Chosen as a normal pnrasit~ of Popillin luadliguttata hut later imrestigations revealed that it would oviposit and develop (qunlly as well on P iaponica The present knowledge of the distribution of T canalis indicates that it covers a Oreater rlmge of territory than was at first supposed ancl it is now known to occur at 7 localities in Chosen 4 in China and] 6 in Japan Although a normal parasite of P quadriguttata in Chos(n it is nssnmed that its normal host in Japan i P japonica since this beetle occurs in all localities where this Tiphia is found A full account of the life
---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
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34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
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_ - shy__ middot_rmiddot
is
j
L
r
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---
INVE8TIGATIO)fS OF PARASITES OF TAPANESE BEETLE 15
history and general hubits of T rernalis nt Slligen Chosen bus been given in previous publications (910) ~ Previous to 1926 nIl shipments of Tiphia rerllalis consisted of cocoons reared nt Swgen The work of propagating this materinl not only involved a tremendous amount of labor in rearing itnd packing but necessitated laboratory overwintering in the United Stutes alld mnting of the emerged femnles the following spring before colonizntion in the beetle-infested nrea could be Ilttempted While studying the life history of T vernalis tlt Suigen it wns found that the femules lived approximately 6 weeks under laboratory conditions In 1926 a trin shipment of adult females was made in speciully constructed shipping containers provided with food and water for the females during transit Although the material was ell route approximately 25 dnys this method of introduction proved sntisfactory and later investigations in the United States showed that imported females lived an avemge of 25 days t1fteL nrrivul and deposited un average of 38 Iggs per femnle (11)
Shipments
The work on Tiphia rernalis nt Suigen Chosen during the period 1929-33 consisted in collecting adult femnles in the field nnd shipping them to the Gnited States In nddition some rearing work wns cRrried on in 1933 Owing to the rapid transformation of Popillia japonica grubs to the pupal stage after the middle of June tl11 cOllsigIlln(nts of adult females were forwarded so ns to arrive at their destination before June 15 In table 5 is R summary of the various shipments of udult femnles from Chosen to the enited States from 1926 to 1933 inclusive
TABLE 5-Summary of shipments of adults of Tiplda emalis from Chosn to the (nited States 1926-33
I Adults I Adults survivingYear Shipment shipped shipment
I ---------1 1--shyNllmbltr l-umber t Peru7IL
990 99 100 1926 bull W~~~middotI 1515 43 f S
1800 i 312 li3 1927 _bullbull _JFirst _bull _ _ a
t 584 2 S-4 t ~ iua
llecond -_ - bull 2 370 ~ 31762S4 t 32lJS I 1396 423
159 40~~~ i 4544 Sli5 5 iOO i 3 r~9 6-1 i lf~middot bullbullbull I 310 ~ 1233 911 560 49-12 I 87 )2899 2736 914 2S19 I 2~2a 895 33m alar t 933 83i9 T JZi 1 RoO
12412 10237 S25~=Jt~~~~middotmiddotbullmiddotmiddot imiddoti Totnl or averagebullbullbullbullbullbullbullbull _bullbull _ _ __ _bullbullbull _bull __ HO257 4i413 6551=====Total or average (or the 5-year period coered h~ this bulletin _ 4iSI5 40193 8-100
It will be seen that with one Ixception tlH first consignments of nduIts each year which wer( collected shortly nft(r (ll1cl-genc( nnci before nny cxtcnsiyc odposition hlld b((n fHcoll1plislwd glle II
higher percentnge of sUlTi ul than those coLL(ct(d lllter in the S(tSOIl
The percentnges of surviynl in shipmr-nts mnde nftN 1928 un notielshynbly highel nnci this is Ilccount(d for by It chl1ng( ill the food In
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
16 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
the earlier shipments the food consisted of a weak sDlution of watel and honey or sugar After 1928 the food consisted of pure honey and pure powdered sugar kneaded into a composition resembling cundy
After the termination of collections for shipment in 1933 females were still abundant in the field at Suigen and it was decided thnt this material be ulted in rearing work From 1lny 24 to June 26 a total of 11031 females were collected from which 65G53 parasitized grubs were obtained These produced 37045 cocoons or 564 percent Of this number 28380 weIe shipped to the United States
Owing to the very successbl establishment of severnl colonies of Tiphia 1ernalis in the beetle-infested arel in the Lnited States from which thousands of females arc nxailnble for colonization each yenr no further introductions of this parasite are neceSSl1lmiddotY
TIPHIA POPILLIAYORA Rohwer
T1phia popilliarora wns first described from specimens collected atmiddot Ioliolm and Koiwai ill northelll Japnn in 1920 The next yeur a few individuals were found at Yokohnma and subltequent iIln~tigations revealed the presence of this species at 16 Iocnlities in Jlpan 2 in Chosen and 2 in China To date surh nnatomicnl diflerences ns hilve been found in examination of specimen mnterinl from these 3 rountries haw not been sufficieIltly constant to wnrrant the assumpshytion that more tilnn 1 species is irlYohed Tlus is ulso the opinion expressed by Allen and Jlynes (2) As is also pointed out by these authors howenl there mc biolng-icnl tliffennccs betw(en the form occurring in Japan and those otrllrring in Chosen and China und because of these (liflNences indiidunls from each country ure refelT(d to us a separnte strain or race T popillialoa is distillCtly u parnsite of Hl( grubs of yurious species of the genus Popillia and dtl)(nltis on the host material ilyailable in tlH countrv where euch strnil1 OCCUIS
In Jnpnn it is pnrnitic on Popilli(L Japo71ica in Chosen on P atlllshycOfrulw and P qlladrigl(itata and in China on P formo8ana P chinemli8 nnd others of the same genus
As discussed in a preyious buU(tin (10) th( times of ndult emNgence in the field of the three strnins in thcir respecti-e countries ar( noti(eshyably diffNent The Jnpanrs( strain in northern Japan eIHIrgcs about the middle of August 01 npproximat(ly 2 w(eks (arlier thnn the Chosenese strnin nithough the monthly m(an tempemturls at Suigen Chosln are higher during thl period from F(brunry to NOHmbN thnn the monthly menn tempemtures at Koiwai Jnpnn At Xnnking China neal whieh the Chin(se stmin is found tlH monthly menn temperutur(s nre highlr throughout the year than at Suigpn but the Chinese strain docs not enwrge until the laU(r part of September 01
3 ieeks later than the Chosenese strain Thus there npP(llr to be distinct biological diflel(nCes in the time of apprarnnce in th( field of the Ulr(e stmins regardless of the nuiation in t~nJ peTfl ture in the three Countlies This 2-week difl(r(nc( in the time of tm(rgellc( between the Japanese and Chosel1Pof stmins also plmiddotenil~ in tite enitN] States wh(l( both stmin hn( bpeIl slIbjpetN[ to tIl( SilJl1e
tlmpcrntures From 1920 to 1923 indwiV( renTing work wus cilllied on at
Koiwni Jnpan with tilr Tnpil1eSr stmin of Tiphia ]JIJpillialOra und extensive shipm(nts of rlaJ(d (o(oons weJ( mudp to thegt lnitcmiddotd
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
IXYEi-iTIGATTOXS 0]0 PAHARITES or TAPAXESE BEETLE 17
States Owing to the establishment of this strain no further shipshyments of it ere lIlade ufter 1923 CXC(pt for two unsuccessful shipshyments of adult females in 192j and 1926 In 1925 1926 and 1927 large shipments of rearcd cocoons of the Chintse strain and a very slUall numbfl of reared cocoons of the ChoselHse strain wcre made
In 19]0 it was felt that the Choselles( strain bfcause of its later nppearance in the field when the grubs of Popillia iaponica arc larger nn(lmole fully den-loped would be more suitablc than tht Japanese strain ill tho l ~Ilited StfLtes H(-aling work was again taken up at Suigrm but owing to n limited supply of f(males in thl field only 979 cocoons WNl (llr(d nnd shipprd in 1930 and 193]
TJIHn ~JTllA Allt11 and luynls
In 1925 Tiphill 1iwtuTa a parasite of seY(ral spring and eally-sumshyn1(1 sp(ci(s of Popillia principally P CU1J1icollis was found in the district SlllloUl1ltling Shillong in thr Khu8i Hills of Assalll Proyince India Th( breeding glOUIlds of this species W(IO locntod in the spars(ly planted pinE forests in WIY hilly country whereas the feeding grolinds were found se-eral mil(s uwuy at the Cl(st of n runge of hills where 11 considerable growth of shrub= of -urious kinds WitS hewily inflstNl with Itphids _8 pointed out in a pre-ious bulletin (10) this wide sepnrntion bptn-en the br((lding und fl(dill~ nrens requiring the femnles to trayel through severul miles of forest for fpecling nnd then to return for o-iposition cnlls for it Y(lT Imusua] nnd inter(sting hnbit with thi~ glllus bull
Extll1si~I rNlring work with Tipkia lItatllm was carried on ut Shillong in 1929 after whirh the work in India wus discontinll((l The sprin~ of 1929 likp thut of 1928 was unuSlwlly wann m1ltL the Jirst indiyiduals w(lr found in the field on April 15 They (lid not become abundant howpnr until n ftp] the spring rains which W(I( latet than in 1928 On April 17 the sex rntio was nbout 5050 ufter which the number of mnles increased so that by )Iay 20 th( sex 11tio was approxishymately 25 males to 1 fmule As u result of deur warm wlather during the Inst 10 days in )Iay thl lfitio of mules and females lH-ailshyable for coll(ctioll aguin becnme nOl1llul This ppllo[L of good weather was followed by another spell of min enrly in June
The collection of flmalls for refl1in~ work was b(gnn on pril 17 find continued until Tune 25 The rains during the llliddle of ~ray and eurly in June s(riously hnndicnppedmiddot collecting und only 7786 f(males were collected as compared with 12606 und 12418 felllales in 1927 und 1928 respectively The lllnximulll numb(l of females in oyiposition at one time wus 2168 as (ompared with 5054 in 1928 The greatest n umber of eggs deposi ted in it single day WitS 771 from 1893 females and the total numbCl of oYipositions obtaillPd during the entire period mostly on grubs of Popillia wpricollis WllS 24845 The general uyerage wus 030 egg per dav pCI f(male as compared with 039 in 1928 v
In rearing the cocoons in the laboratory a medium of soil llnd chopped moss was used instead of the usual composition of soil and sod This combination improved the physical condi tion of the soil and retnrcled excessive loss of moisture thus increasing the rate of cocoon fonnation From the 24845 grubs parasitized 8012 cocoons were obtained or 323 percent
234374-40-3
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
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CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
18 TECHNICAL BULLETJX 738 U ~ ])f~PT OJ AGlUUULTUlm
Shipments of liphia maiula cocoons in previolls years did not leave india until the fali but in 1929 tll( cocoons were divided into 2 lots and fOlward(d to the rnitNI Statps in th( Slimmer immediutelv nltor they (1e spun Thl first consignment comprising 1800 cocoons loft Calcutta India July 18 und aniyed at thl ~Joor(stown laborashytOly 38 days lat(r 111( mn tfrial was placed on the bont at a tlmpershyatur( of 45deg F with till undllstanding that this temJ(lature wouLd be maintained throughout the yoyugf rlt( uwthod of packing the cocoons differed sOlllcwlw t from thnt ImploT((L in Inpall -ooden tmys 15 inches squall and 2 illciIPs ([(ep poundittN] with cross-section sets plOyided compurtm(nts j inch SqlUUC and 2 incIPS delp Two cocoons (ach surrounded with moss and sepamtc(L from (nch other by a sQUl1l( of tin 01 cdluloid well plnced in each compartment Th( trays ench holding 450 cocoons w(re tlHn pack(d in large WOOdlll CttS(l in Slts of fin surloundlC1 hy a InYlr of fresh sphngnum moss
Thl second lot of 6000 cocoons wus shipPtd from Cnlcutta August 8 and wus Ill route 38 (lays atmiddot a temp(raturc of 52deg F 10 simplify the parking and leduc( tlll handling at th(ir d(stilHLtion the cocoons in this shipment w(1e pnckCcl incliyidllally in -iuch yials stoPP(1(d with ubsorblnt cotton all(1 thls( yials WI1( ph1ltrd in llctanglllar trays of perforat(d tin tUld hdd in plac( by cllrclboard cross sections The trays (ach holding 200 yials and sl1llOundrd by n 2-inch lavCr of fresh middotsphag-num 1Il0S~ W(I( pack(d ill ll11g-e woodell raslS
T11( c01ulition of the (ocoons in titese two shipment wns CxCCpshytionally g-ood on 1I1i11 ut ~loor(town X r as only 16 cocoons showed allY signs of lxt(lllnl funglls
TJ[1Il PlLIIVOH Allpll and Jaym
During- the ilwestigtttiol1 in India n small 5[1(gtcie of Ti phia I1tN describ(d a Tipltia ]17dliloa as found in the Khasi Hills of Assl1m P1O~ince This sp(cits is found in Iarg(gt numhrrs ill the field from the lattrr part of July to early in OctobCI Althollgh it prillcipal host uncl(l normal lilllt[ conditioll is a smnll undett11ninpd nlhline b(ltll it will also attnck and d(plop 011 ~prOI1(I-insttn and (urIy third-instal grllh of Pllpilia cupricollis and nnother sp(eips o[ PO]lillia Because of this it wtts thought tltut this s[Jp(ies migh t bt of [0111( yalue as a parnsitp of P japollica in the summpl and early part of the fall whcn thl grubs are small
Previous innstigation hopd thnt rCiring work with this sprcits was UlUlPCeSSal owing to tht hltglt llumlwr of co(oons thnt could be collectpd in tll( fideL Tn tllP flll of 1929 a total of 21OO= co(oons were colhct((l and storp(l in it (001 place iwaiting shiprrlPnt thl followshying spring Innstigntiolls hOI (1 show(d thnt adults from shipshyments of this s])(ci(s did not (nl(lgp at a time snitablp fol liSP against Popillia japonica in tht (nit(d States and instructions WNt spnt from )fool(stown X 1 to cliscontinu( all work with Tiptia plLllirora and not to ship uny cocoons on hand
TIPI1A srgt Taiwan J
At Hokuto n(al Taihoku in north(ln Taiwan adults of an und(shytermined species of Tiphia WI)r( found in the field in 1029 from the latter part of August to (ady in October This spcci(s o-ipositcd
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
DESTIGATIO~S OF PARASITES OF JAPANESE BEETLE 19
and d()n~lopecl very readily on grubs of a species of Popillia nnd it i~ assnmed thl1t the nomlal host in the field is a member of this genus The adults feed at the nectar glands found at the base of the leaf petiole of the sweetpotato and other plants
ThC rgg which is dark gmy is placed Yontrnlly between the eighth and ninth abdominal segments with the anterior pole directed out (fig 1) During the h1tter part of August and early part of September thl Cgg stage coyers 4 clays tlllCi the larval stages apprmimatcly 10 days
Because of the scarcity of this species in the field in 1929 only fL
few females were collectCt and from these 911 parasitiz(~d grubs were obtained These produced 387 COCOOllS a percentage of~425 Of these 379 WlrCmiddot shipped to the United States
TUE PARASITES OF RELATED SCARABAEIDAE
During leClnt vealS three additional scarabaeid beetles from the Orirnt lluH become estilblislHd in the eiLstlrn part of the rnited Statls Thl oriental beetlc (Anomala orientalis Yaterh) which ns first found in Connecticut in 1920 has spread to ~Iussfichusetts K ew York and X ew Jersey find the Asiaticgarden bCetle (AlltoSelica castanea (A1lOwraquo) which WiLS originally found in N ew Jersey in 1920 is now known to occur in Iassachusctts Connecticut X ow York PeJ1l1sylYtulia Debwarl Maryland the District of Columbia and South Carolina SellCa peregrina Chapin which was first found on Long Island X Y in 1927 is not known to occur in allY other State Owing to fayomble bree(ling conditions these pests 11111 inclrased and spread causing considerable damage to truck crops OlW1l11(tltal plants nnd Inls especially in suburban areas which arc particularly suitable to both adult and lan-Ill feeding
Ol1tsicle the enitfcl StatCs Awmala orientalis is known to occur in the Hnwniinn and Pbilippine Islands nnd ill JlLpan In Tapnn so fnr as is known theI( is but OIll genCrntion il YClU the adults lwillg pres(nt in the I1eld from tlll lattl part of Junc to carly in SlptcJIlI)(l The ndl1lts which ar both diurnal nnd nocturnnl in habit pxhibit little nctiyity in the lolnnds during the day but tlwre is consilblablp dnytitlH nctiyity- in tll( mountainous rCgioI1s Although this bppte is fairly common in some sections of Japnn there an llO IIcords in Tnpnl1tsl littrntulC QI its b(ing of any lcol1omic importft1Hl to agrishycultural crops and the writrlS han nCn[ obsery((i it f(lding to any extCnt on any plants of importnnct Iost of thl adult f(((ling is confill(d to yurioHs sptcis of PolYfJon1l1n find othll hllbuctolls pli11ts which ul( conunon throughout Jupfin
Auto8etica castanea occurs in Japan as w(ll ns on the Asiatic mainshyland fn Japan it is tlH most common of ftbout 35 SI)lcies Iwlonging to til( Sllicina0 Tlwrp is only 1 gP]1(Imiddot(ttioll il Ylar ti1( adults app(lrshying in thc fidel from parly in TUlle to SCptrmber TIH adults bring nocturnnl iLl( mrdy s(en during the day but on warm still nights tlHY nrC abundant in some sCctions A fayoritc piuc fol th(tl1 to (ongreg-atr at nigh t is around lights As 11 l)(st of (anomie im porshytance in Inpnn it is of little consCqurnce Soml dumagt hus b((ll rrportCd from the feeding of adults 011 young Jl11IS(IY stoek nnd omnmCntnls find from grubs fceding in lawns YIH1l grubs of this
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
20 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
species w(re bping dug for parllsite-lenring work nt Yokohnmn they were found to be most abundnnt in rather dry soil around the roots of Artemisia spp and other weeds bull
Serica peregrina is known to occur in Inpnll Chosen nnd China It has only one generation a year in Japan the ndults being present in the field from late in June to late in August Although this beetle is not considered a pest of mnjor importance in Japnn nnd ChOSN1 the adult is reporttd to feed at times on apple penr mulberry barley cauliflower and aspnrngus in Japan and all millet cotton and sugar beet in Chosen (8)
Although these beetks nre considered of little economic importnllco in the Orient their spread mel lllcr(asing importfl11c( as pests in the eastern part of the Fllited Stntes hns lHc(ssitatec the illYCstiglltioll and study of their parasites in their native homes
During invcstigntiol1s in Japnn Chosell and Taiwt11 no sllitubll )arasites of the adults of these thTee b(etlts w(re found J[alwria incongrua which is a parasit( of both Popillia iapOllica and l71ulilala orientalis in JapUJ1 has three genrrations a year filld rrquins fill alternate host to complete its YNtrly li[( cycle Ko fittempts 11(1(
b(en made to introducr this tachinid pnlflsit( against A oripntali flS npeatecl efforts to (stablish it agflinst P japonica in tl)( rnihmiddotd States have hiitd In Chosen an lllldetclmined tachinid was fonnd which is palflsitic on a spNils of Sericcl hut this parasite apP(alS in t11( fie1c1 during lay which is too carl)- to 1)( of all) vfilup ngninst A~ltoserica castaneamiddot or 8 peregrina in the presellt UlPI1 of inf(sbltion
PARASl1ES OF THE LJlYA
SCOLIIDAE
~Inny SI)(cies of Tiphia parasitic 011 Anomala illl(l 8(rica W(fl
stultiipcl in Japan C110S(11 nnd Tniwnn but only 11 11( consiti(Ild of sufficicllt impo1tnne( to warrant illtlOeildion 1thollgh SOJ1H of tll(S in thrir nativc hahitnt arc parasitic on grubs othpl than thos lstablisJpd in thp lnit((l States inv(stigntions htlYl shoWIl that the Rp(eirs of Tiphia arc in gelHlnl g(IlNir ratiIpr thfin sp(eiJk in tlwir (bojc( of hosts Thrse specics of Tiphia mo1(OC1 ill( locflliz((Lin thcir distri~ution and aTC dependent on suflki(nt supply of udult food and sUltl1ble host grubs
During th( period from 1929 to 1933 the species list(d in tab( (j were ilwestigntNl and 77270 cocoons and 6139 ndults w(rc spnt to til( United Stutes
INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
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34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
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INVESTIGATIONS OF PARASITES OF TAPANESE BEETLE 21
TABLE 6-Species of Ti1Jhia parasitic on scarabaeid grubs of the jenera Anomala Autoserica Phyllopertila and Serica investigated in Japan Chosen Taiwan and China and shipped to the United States
Spltlcics oC Tiphia Countries HostsI T aericae Allen and Jnynes-------ITfiPan Chosen China __ _1 Serica Spp1
bmiddot 111at 11 d J J IAllomala 8cholleldU Ohnus1
AnOmala cltprea Hope bull bulllIomala tesfaclipes Mots
T Isec a Cn an ayucs bullbull -- -- apan _______ _bullbullbull _ ___ _ tuomulu oriontali bull
T bltrrelll Parkerbullbull__ bull___ bullbull ____bullbull___dobullbull _____ Allomula 8chonfedtiI I
Anomala rltoClLprta middot k d Serka Sp1T castantaerora 1 r ermiddot __ ~ _ ~ __ middot~_Imiddot-- __ Omiddot~___ _~ ____ _middot _____IAulo8crica (a8tanea
middot P k J CI Serica Sp1T homonc111arra ar cr___ _ _ ~_~_ apRIl IOCD ____ ______ Autoserica castanea
m bull I I P k J Serica Sp~ 180 a a ar er_ ___ _ ~ -_ _ _ tlplln_middot_~ ____ bull____ ______ l bull411toserica caJtallea
bull I IPhYIlOJlerlha cOlIspllrcaa ITnr1 I _ c i Phyllolerfha pallitlipcllis Reitter
T lotopota alienI Roberts -----middot1 Japlll Choen lunn_ Anomnla orieltalis I I AIIDl1Ialu schollfeltlti
7 saoi Parker _______ _ Chosen_ _______bull_____ r SeriC( Spp1
1 sUmla Parkcr----- _ --l Jnprm -bull--- ---- - -----~e~~c~(~l~P~honfeltlti T I III I J p i d i1110 maiu rujocllprea 1
Iegltp aua en ltlD( nncs___ - omiddot----middot------middot --i AllolUal orielltalis I t PAylioperlh conpurcata
An umdentIfied speclcs __ _______ _ lnlwttn----------- bull --- Anomal SIl j i
I From field obsermtioIJ
TIIHlA ASERrcAE Allen and Jaynes
Tiphia ascricae a moclrratr-sizrcl sp(ci(s wns first found at Suigen Chos(n in 1924 and has since b((n I((old(cl from China and Jnpnn In the Chosrn arCa whrIr it has a l-yrar life cycle it is very abundnnt nnd is pnrasitic on various species of seliciIll grubs during June Ilnd early in July Its life histOly hns been fully tlcnted in a previous publication (10)
During tIi(gt years 1929 to 1932 inclusive 1672134 oYlpositions wCIe obtained in the laboratOlY giving the comparnti vdy high rate of egg procllletion of 097 egg pel day per female From these oviposishytions 721306 larval or 434 percent dcn-ioped cocoons a much hightgtl rate of cocoon dcrlopmeut than that obtainrd in the brrcding work cnnied on in prfyiolls yrals During this 4-yeal pfriod 131270 of these (o(oons were shipped to thr Fnited States Early in Junr 1929 11 consignm(I1t of 1656 adult femairs was forwardNl Although the mtthods of paeking nnd thf food used w(rf the sume~flS in tIllt SllCCCSSshyfui shipments of Tiphia lelllalis only a few adults W(l( alie on arrivnl at thrir drstination The results of txpelimcnts cflrrircl on at Snigrn in 1929 to clfterminf the longevity of adult femalts whrn storNl in a cool pincr undrr shipping-can conditions jndica~((l that thr length of life of this spcci(s is considentbly shortfI thun thnt of other mrmbrls of the grllus w-hich hUNe hef1l workrd rith to dnt(gt and that thr high mortality during shipment is due to this faetor rntlwr than to fuulty shipping methods
TIPHIA BISECUIATA Allen and Jaynls
Tiphia biseculata is normally parasitic on Anomala spo but il1Y(stishygation s11Owrlt1 that it would drvriop readily on Popillia japomca in thr laboratory ConsrquCl1tiy it was imported into thr Fnitrd States from 1924 to 1928 inclusivr flS a parasite of P j(Lponica Since
22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
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34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
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22 TECHXICAL BULLETIX i38 U S DElT OF AGRICULTURE
investigations in the United Statrs showd that this Tiphia was of littlr valur as a parasite of th JapaJl(se bCet1r in the present arCas of infestation JlO further work was done with it nntil1932 At this timr a shipment of cocoons wns macIC from whiCh the emerging femal(s w(re libCratrd as palasitrs of AnolnaZa orientalis which was becoming of increasing importance as a pest in Lhc vicinity of New Y01k
This Tiphia is one of th( two speeies which haY( two distinct genshyrations a yat in Jitpan Th first brood rppars in the field during June and those of tIl( s(cond brood from late in August to lat in Octobrt TIl( first brood is much mor( numerous than tIl( sccond
Previous to 1932 this species had bcn found in only on locality in Japan at ~l1ho a small sandy pninsula jutting out a f(w mils into th sa llar Shimizu in Sbizuoka-kn In 1932 Tiphia biseculata was found at Yokohama and Chigasnki Jnpan till last-named locality having much the same Cnvironm(nbll conditions as those found nt ~fiho
Experimen t9 currird on with thr first grnp1l1tion in 1932 show(l that this speci(s acc(pttd grubs of Anoli1aicL olienialis tq tlully ns readily flS those of A schoenjefdti it3 normal host Rrilling work was cilrried on with th( srcond grntrntion nt ~liho from August 23 to OctobrI 24 1932 nnd during this timr 3649 fnHtlrs w[ collct(1 Femalts w(r still abundant in th fi(ld wlHn collrction ((nsNI A total of 38919 ovipositions Wll( obtained whieh protiucrd 8507 (o(oons or 2184 petcnt of which 616 or 158 pltlcent of th( totnl ovipositions wre nff(ct(l by fungus This smull pltC(n tagp of co(oon formation which is low ns compl1llCl with many Sl)((i(s of Tiphia is a(countrd for by tilr filct that tIl pnrnsitizrd host grubs move about l great denl in thr oviposition tins and r(llling Hats whieh tnds to dislodg thr eggs ilnd young 111ln In tIl( fnll 7692 of thts ~ocoons n~rc shippd to the Lnitrd Statls
T[PHIA RLRREILI ParkN
Tip]Jia bU7Ielli was filSt found nt Kngnmignhl1rl Jnpan in 1931 and latrr wns found in small I1umbrrs at ~Iiho and Hisnznki on the island of Honshu find at Ishiki on til island of Shikoku Its normill hosts are tl large grubs of Anomala Cllprea find A lestaceiZl(s but it is palilsitic also on A 8cholljeldti ilnd A l1~fOC1lplecL Sincr no glubs of Anomala orientalis Wll a~nilabhl lnbollltory tlsts with this spcci(s Wlre not att(mptNl
Tll adults art found in the fillt1 from til first of Sptll1bpr to thn middle of Oetolwr f(eding on tIllt hOlltydtw of aphids nnd otlH1 insrcts TIHI is on g(lwrntion II yt111 ilnd tht~ wint(1 is pnsscd as th( mntult lnrnl within t11 (o(~oon
Tll gg which is grnyish whitl is plnc(d nntrally bptw(tl1 lh last thoJflcic aTHI first nblt1ominal sgmll1l (fig 1) rndIS(ptltgtmbf1 templilturps at~Iillo th incubation 1)(lioltl is dnys and th( Inrntl f((ling pliod nPflrly a mOllth flip Inlgp cocoon is imilnt 10 thnt of Tipltia brerilineata in having n thick fltitlik (oPling pad with illl UIH(n smfaCr Tht (ocoon is n dnrkpr brown than thosp of most Spt(ilS of Tiphia Ilnd sOll1rwlHlt glnZ(middotd lnekinp nImost ntirlly the 100s( ontl stIflnds of ~ilk
Owing to tht H(flLcity of ndulti only 277 Oip()~itimH Wlrl oiJtnillld in 19i2 of which (H d(gtYllopld to the cocoon tngl ~iillg II d(nlopshy
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
IXn~5TlGATIOxs OF PARASITES OF JAPANESE BEETLE 23
mcnt of 22 percent These cocoons were shipped to the United States for colonization aguinst Anomala orientalis
TIPHlA (tS1ANEAEVORA Parker
Tiphia castaneaelora hith dostly ltstmblts T aSeliCH6 taxonomishycally WHS first found at Nalashillo near Yokohama Japall in 1931 and the next fcmiddotar it was found at Tachikawa neur Tokyo In tht1
field it is parasitic onlericinE gmbs and in the laboratory it was found to oviposit nnd develop vcry lNulily Oll hltoserica castallel~
The adults mny be found in the field from the middle of May to the ll1(l of June (middoteding in lHuubels on the blossoms of chestnut find oak There is a single gPllcration (1(h year and the winter is passed in the mature larval stage within the cocoon
Ttl( grayish cgg is placed in tht sutUI( bctw(en the fourth Ilnd fifth stgments of tht grub just vclltrnd of the lateml line and with tllt an terior pole directed toward it (fig 1) Tlw incubation period lasts from 3 to 5 dnys and till denlopment of tllf larva to the spinning of the cocoon rrquirrs 10 to 12 days Thc co(oon is small and light brown with It modrmte filament
From 1153 fcmalrs collpctrd at Tacbikawl) in 193i n total of 9436 oyipositions were obtninlltl giving an cgg-production rute of 085 pCI day pel femnle Thesr oipositiolls yielded 5a90 cocoons or 571 prrcent cocoon formation Of thlse 5066 werp shipped to the rnited States for (olonization
TIPHIA 1I0(OX(tLARI8 Parker
Tiph-ia homonc1tlctri 11as been found in five localities in central Japan and at Suigen Chosen It is paIilsitie on scricinc grubs und under laboratory rcaring conditions develops very readily on AUtoserica castanea
The adults hosp food is thr honeydew of aphids and other bugs are found in thc firld at Yokohama front tiJr first of Septrmber to the first of October There is one generation it yeitr and hibernation takes place as a mature larVl) within the coeoon
The eggs which are slightly gmy are placed n~ntrully usually between the third and fourth abdominal segments on tbird-instUl host larvne and usually one or two segments nearer the thoraX on second-instal grubs with the nnt(rior poip directed outwnrd Apparshyently sccond-instur ilnd thinl-illstllr grubs 1re equally favored judgshying from the Ilum belS of eggs in e[tch of these positions on both fieldshycollected nnci Inbomtoly-plllUsitizeci grubs Tile incubation period lasts from 3 to 5 days (Ie pending on temperatures and the deeopshyrnent of thr lana to the spinning of the cocoon requires approximately 20 days in Heptembelmiddot Tbe cocoon ii rtltiler smllll but othenvisr norshymal in form nnd eolor
This spe(ies apparently ha a 11igh n1tr of oviposition as 6G femnlrs in one experinlPn t deposited nn fLeragr of 111 egg pel day pel female Although this species was moderntel~ Ilbundant itt Yokohl1lna in 19~2 only lirllited rrllring work was carried on owing to difficulty in ohtainshying It suHicient quantity of serieine grubs From 1101 ovipositions 274 or 249 prlc(nt (O(OOIli were obtnined of whicb 10 W(IP ltflecmiddotted hy fungus The 264 hrulthy (ocoons werr shipped to the 1~lJitld Stutes il 1932
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
24 TECHNICAL BULLETIN 738 U S DEPT OF AGlUClLTURE
TIlHI ISOLATA Parker
liphia isolata 11 smu1l species pall1sitic on sericine glllb5 wus first found ut Yokohamu uud luter ut Toyosu and RokkoSllll on the islnnd of Honshu md at fhre loculities on the island of KYllShll Ll1bllshyratory investigntions showed that it oviposits and develops readily on Auto8erica castanea
In the 7okohama district the llc111lts which feed on honeydew are found in thE field from the enrly part of September to the middle of Odober There is only one generation a yeal and hibernation occurs I1S the mature bryu within thl cocoon
The pgg which is placed with occitsional variation yentraIly l1(1tr the medillll linp tll1d between the selond Hnd third tholtlcic segments (fig 1) is llearl~T whitp when first deposited but dnrkfns with nge Till anterior pole is dileclrd outwHrd The incuhntion ptriod covers from 4 to 5 dn)Ts nnd thr development of tilt InlTnl stnge requiles from 20 to 25 dnys The cocoon is normnl in (0101 but raquomnU with no fluffy outpr (oYering
Owing to thr lInnll nUll1brr of sericill( gruhs nynilnhle extellsivf (nlings were not possible nnd in ] 932 only 79 (o(00n8 were obtained for shipn1ltn t to the rnited Stlltes
fI PH IA NOTOPOIITA ULENI Hobert
Tiphia not(lpilifa allPni WitS found in 1922 lit Suigln Chos(n whEI( it is primarily it pnm8ite of Phyllop(ltw COllspuIcata although somE indi-idunls nttn(k P pallidipuznis In 11)24 it W1s found itt Prnniu Hangrhow and Kuliang Chinn
In 1931 nnd 1932 it wns fOllnd nt s(-eml wiltirly s(pllllltfd lo(nlities on Honshu Islnnd Tnpnn At Xngnno nnd Toyou in ((ntml Jltpnn ther( is (-idEIHl to dnt( of n plItinl se(ond g(n(Intion similnr to tlllt of 1iphi(l tgitiplafa os somr in(iyiltiunls npP(HIlnt( in Jun( nnd EuriS in Jnly tilltil prng(ny (1l1(lging in Rlptrmber This spl(i(s is mu(h more nbundnnt IntI in thr silmm(I nnd cnrly in til( fnU howenI appenring from litte in August to lilt( in S(ptEmlHr Although no inycstigntions wel( (itlrifd 011 Itt Yokohnmn Illlcl ~[iho with r(f(rclI(( to the first gcnemtion it is lH(U111Cd thnt th( snmc s(Hsonnl eydc preynils in these IOClliti(s At ~riho nnd Yokolwmo thl ndults ItImiddotp found ill thc field from the Iattel pnrt of Augwn to til( (tIdy purt of Odobel At Nilgnno th( (uriy adults f((( on til( hon(~middotd(w of uphids on soyb(l1ns nllt on thc hlosoms of Pol1f[Jolllm reynolltria while til lute illdi~idunls [(cd 011 Allium orommmiddot At ~rib() they feNl on the nectnl glnnd nt th( bnse of the lenf petiolI of sW(ltp()tn to und nt Yokohmnn th(v fe(d on white um b(llif(lOlIS hlossoms
At X nguno ulHi ieinity Phylloprtlta (OTl~p1lfc(If(t npP(flrs from fi(ld StllHYS to be th( normnl host In til Inboll1tory howrv(I it rpndilT pfllllsitizfs Anomala orilntais though co(oon formntioll on thi host WfiS POOl At Yokohnmll pnlnsitizNI grubs of A orinliali -1 sctOIlshyfeldfi nnd Phylloper-lha sp (1( found in the ficld nnd ~Iuhs of th(s( speri(s wer( nlso USN ill th( I(Hring work S((onrl-instnr gruhs of A swnfelrlti were more tcllclily lHctptrd in th( Inborn tory work thtlll third-instnl grubs d the fHlIIH sp((ifs
Th( egg wbieh is lght grny wil(11 first lnid hut turns darkN b(fol( hut(hing is pllc(c hetw((n the sc-enth find righth rthclomillltl stgshy
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
INYESTIGATIONS OF PARASITES OF JAPANESE BEETLE 25
ments ill the Jepressed tHeft near the hlteml margin with the anterior pole directed outward (fig 1) The dumtion of the egg sta~e is 4 to 5 ltlays and thn t of the larval stages 15 to 17 days under September tempemtures The winter is pussed us a full-grown lnrva within the ltocoon The cocoon is of medium size loose and fluffy and not sepshyarated by distinct layers
In the fnIl of 1932 rearing work wns cHrried on at ~Iiho Yokohumu and N uguno At ~fiho from l7G femules fi43 parasitized grubs w(re obtained which developed 149 cocoons or 2318 p(fcent At Yokoshyhama 609 females pumsitized 3648 grubs fOm which 526 cocoons or 1442 pelc(nt w(t( obtain((l At Nngal1o poundOm 709 f(mules u totul of 5593 purusitized gltlbs were obtained which developed 919 or 164 perc(nt (ocoons Of ti(se cocoons 133 were shipped to the 1n1ted States th( (ll1ltrging adults to be used ngainst Llnomala orirnshytalis
TIPHIA fAfOI Parker
In a predous publication (iO) Tiphia maZalana Cfim (7) was rf(ll(cl to us occurring in China find (hos(n but at that tim( it was thought that two distinct sp(ci(s might be involv(d Oll nceount of mnrk(d clifr(t(nc(s in life history and hn bits In 1932 a sp((i(s of Tiphia wus found in Jtlpnn which at first was thought to b(gt T malayarw und bioiogiCal studi(s w(re (onetuctNl on both th( Choscnshyes(gt and Japanese forms In til( cours( of th(sc studirs biological diff(r(nC(s nrp obs(1((1 whieh prompted fi cios( inyesti~a tion of th(ir taxonomic chnlllet(rs Thjs r(sultrd in tlw drt(rninntion of the ChoseTl(se and Japanrs( forms us two distinct sprei(s und these wel( lat(r drs(ribed as T satoi from ChOS(ll und T sternata from Japan Comparisons of the t vo sp(cirs w(r( thcn mtldr with sprCishymNlS from Chillll d(signu tNl us T ma[alana und these rev(uled tuxonomic diflerenc(s by which mMplinl from tlw thrre l(gions (ould be s(purnt(d Fulth(r study shorNI that non( of the thl(e sp(Ci(s in qu(stion wpr( T malaana und the form hom Chinu wus later cleseri beel us T frater (18)
Tiphia satoi has b~n found in fiv( 10caliti(s in cennal and soutll(lI1 Chosen hut I1rY(r in finy gr(at nbundunc( except at Suigltn wh(l( it wus originully found It is parasitic on the grubs of yurious s(rieine sp(ci( thut arc in th( third lnnfil instal during the spring
The udults appNlr in tlH firM from th( middIr of April to thc (arly part of JUll( this bCing the first sp(cirs of th( grn us to appNlr in the spring in Chos(n For the most purt th(gt adults f(pel on the honCvd(w of aphids find o(Casionally on the blossoms of F07sythia sp So far as is known th(rp is only one gCl1(gtrntion n Y(IlI unci till wintCr is passrd as Ow adult plobably in til( cocoon ThC rgg whemiddotn first laid is whit( but dark(gtns bltpoundo1( hatemiddothing It is p1acr( yrnilully h(twNll tbl tlllrd and fourth abdominal spgmrnts with tbr unlrlior pO( dir(gtct(( outshyward (fig J) TIl( Cgg stngr lPquilps 10 to 11 days ancl tbr lurvn stag(s 20 to 22 days in ~ray Th( (o(oon is of an(utlmiddotnl (0101 and thr olltrl co(ring is 1005r and not sCPlulltNI into IflycrB
In 1932 long(vity (xp(limrnts undpl Bhipping-Call conditions YCrc curried on to ltl(t(Imin( wlHtlwr 01 not adults of this sp((ils could he su(crssfully shippecl to th(l enitpd RUltfS ft(II blting ronfinpd for j w((ks 714 plrc(ll t of thr fpmalrs wprp still uli Tr Ilnd t hrsc lind an
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
26 TECHXICAL BULLETIN 738 1] S DEPT OF AGRICULTURE
avpragl of 2 wleks longlr during which timl thlY wlrl confinld with host grubs and ovipositld regularly palUsitizing tm avelUgl of nlarly 10 grubs plr female
In 1981 86 cocoons und in 1933 46 flnutlps were shippld Of the lattIL 30 were alive on arrival in the United States
TIPHlA qlERNAlA Parker
Tiphia sterTlata a pfllusitl of 81ricil1p ~ll1bs is ttlxonomically close to T satoi It was first fonnd in the vicinity of Yokohoma in 1931 and since then it has blpn fonnd at nine localitips in the centlUI and eastshycpntral parts of Honshu and in seven locnlities in the nortllErn purt of Kyushu
Both slxes npopar in lllllul)(rs from Iurly in Spptember to late in OctobtI The flmale is bPlilvNl to hihprnate undfr ground and during this tinl( tIll fggs take form within the ovarips TIll males appaIpntly do not survi-c hibllI1ation TllEsP suppositions are based on tll( results of laboratory Ixpprinwntation and the fact that to date fipld observa tions hnye shown tIll males to bl vlry scnr(1 in thp spring Females are again ahnndant in the fipId from early in ~[ay to the middll of June willn parasitization of the host takes placf
Eperiments ~with fall-emerged and mated females in the laboratory luwe resulted in a flwisolated cases of oviposition but all other feshymales dissected showed no signs of egg devllopment within the ovaries Periodical dissection of laborntory-hibernatlcl females showed gruclually increasing evi(1en(r of egg formation Springshycollected females oviposit readily under laboratory conditions As has been sniel indications arc tha t the mnles do not SUlvi Te hibernashytion tllPrefore it mny be supposed that Huiles appearing in the spring nre the result of occasional fan ovipositions and serve to fertilize the spring-appenring females It is not known whetlHr this complex ests in N agnno and central Japl1n wheIC the mean temperatures arp lower than those prCa iling a t Yokohama
The egg which is pule gra~T is placld ventrnlly between the third thorucic and first abdominal segments vith the tnterior pole directed outward (fig 1) The egg stage in the spring at Yokohnma lasts 4 days and the larval period 10 to 12 clays In the few instances of full oTiposition that have beCn ob~eryed the egg stage lasted 4 to 5 duys ami the larval stage 20 to 22 days depending on the temperature The cocoon is normal in color with a small amount of (lxterior fluffy filament
In 1931 a total of 200 cocoons WelC reared at Yokohamn from springshyappearing females and shioprd to til( Fnitrcl Stutes In the spring of 1933 2 shipments totalling 4437 females were collertNI from the Yokohama-Tokyo distri(t and shipped to the rnited States Of these 3615 arrived nt their destinntion nJivC
PHfA rEGlTI[AGA AII(1 and layne
Tiphia tegitiplaga which was originaJly found at ~fiho Japan was later found at nine otll( localities in Honshu one ill Hokknido and four in the southern part of Kyushu
At ~[iho tlwre UTe two generations it yellr thl fir3t appearing from late in TunC to tll( lattCl lli1lf of July and the faU generntion from late
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
27 INVESTIGATIONS OF PARASITES OF JAPANBSB 13EE1LE
in August to the middle of September In N ogano in ccntrnl Jfipall two genemtions also occur although lndividufils of the full generation appear to be the more abundant Here the time of field nppefirance is similar to that at Miho The appearance of two generations fit N fignno which is much farther north fincl has a much heavier soil seems somewhat unusufil These conditions fiTe probnbly oftset by the fact that N fignno is in a lfirge valley where the summer temshyperatures are excessively high In both locfilities hibernation of the fall generation occurs in tii~ larval stfige
The spring generation at Mitl~ feeds on the honeydew excreted by aphids on melon vines beans and truck crops while the full generashytion feeds on the nectar glandR at the base of the baf petiole of sweetshypOtfitO find the blossoms of buckwheilL Owing to the lack of truck crops find the sweetpotato at Nfigano feeding of the first generntion IS apPfirently confined to the honeydew found on a species of Liyustlum find those of the second generation on honeydew on the leaves of mulberry
At ~middotliho the normal host is Anomala 8chonfeldti filthough A orientalis Phyllopeltha conspulcaia ond a few sericine grubs were also parasitized A 71foc111J1ea wos the host most abllndfintly parashysitized in the fall find the parasite shQwed a decided preference for second-instal lurnlO Yitll grubs of A orientalis as host materifil the preference was also for the second-instfir lorvae The parashysitization of second- and third-instal lfirvae of Ph11llopeltha COllspurcata WfiS equfilly divided wherefis of the few serieine grubs paltsitized fill were in the third instal In the field at X aguIlo some Popillia Japon ica grubs were found bearing the dnrk egg scar which charncterizes Tiphia oyiposition and in the proper position for pfirnRitizntion by T tenitiplaga (fig 1) Although this Tiphia accepted P iaponira readil~Y in the laboratorY the percen tuge of cocoon formation was exceedingly loW
The pale-gray egg is laid dorsallv on the second thorflcic segment USllfilly near and sometimes on the medifil1 line the llnterior pole directed outvard The duration of the egg stage is 4 to 5 daYR find the Iarvol stage 18 days for efich generntioll The femfiles laid 092 egg per day in the laboratory
Rearulg work was carried on with the ffill genemtion at both ~Iiho and Nagano in 1932 A total of 2020 ovipositions were secured at N figfino but only 84 laryue developed to the cocoon stnge At ~[jho i363 ovipositions were obtained find from these 136 cocoons developed A total of 1180 cocoons were shipped to the lnited Stutes in the full of 1932
TIPllfA 8p Taiwan 2
An unidentified species of Tiphia was conected in very limited numbers at Hokuto ncar Taihoku in northern Taiwan late in the summer and early in the fall of ] 929 This Tiphia the adults of which feed on the nectar glands associated with tIll sweetpotato are present in the field from Inte in August to early in October
The pale-gray egg is placecl ventrally between the fourth find fifth nbdominal segments with the anterior pole directed outwllrd (fig 1) The egg hfitches in 4 days and the lfirva stnge eo-ers a period of 12 to 14 days in September
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
28 TECHNICAL BULLETIN 738 U S DEPT OF AGRICULTURE
Owing to the scarcity of this Tiphia in the field only a few females were collected from which 155 ovipositions were obtained in 1929 These developed 47 cocoons (303 percent) Of these 39 were shipped to the United States
PARASITES ON WHICH NO INVESTIGATIONS WERE CARRIED ON DURING THE PERIOD 1929-33
During the course of investigations of the parasites of Popillia japonica and related Scarabaeidae in the Orient from 1920 to 1928 inclusive the parasites listed in table 7 which were studied and imported into the United States were for various reasons not inshycluded in the work curried on from 1929 to 1933 inclusive
TABLE 7-Species of parasites of Popillia japonica and related Scarabaeidae studied in Japan Chosen China and India during the period 1920-28 and imported into the United States but not included in the work during the period 1929-33
Species Countries IIost~
---------------------------------__-shy-frnchinidat
Pexolllyia venalis Aldrichbullbullbullbullbull Japan__ Popiliajllponiea
Scoliidae JPhyllophaoa lio1llphaliu Datesshy
Campsol1ler~ annlllllia (Fnhrimiddot Japan Chosen Chinn IAUOm((la spcius) Popillia spp
T bull h 1 S th d Serim ariellan Iots rp fa aUfS ml ~ ----~ ~---- --- o -~- --~--~-------- Serica sp
Tiphia 88amellis Allen and Indln~~ Popilia cI~prieolliJnynes
Tiphia biearinata Campron~ Ch AnOmala iereri Heyd)Jnpan Chosen na Phylloperlhu plbicoli Waterh
Allomuu sieter) Tiphia breviliTlcata Aller nnd Jnpan Chosen Phylloperlhu pulJiciJlIiIJaynes Popilllllllwlculf Tiphia duuuni Allen and Jaynes [ndin PopUlia Sp1
MlTilUba trichophoTlLS Fnlrm Tiphia communis Allen wd Chinn~__ A lInericu spJaynes Popilli forllloana
iiphiafrater Parker do~ We~i~~~I~Pl AnOIlUi a 8iertrlfil
Tiphia koreona Rohwer Chosen~_ bull Popillia airocoerllita PhylloptTlha pubicolliI 7iphia uotopolia Allen and (hina~ POJlillia ehineu~
Jaynes Popilliaforlllosanal Tiphia otdorsalis Allen and Chosen ~ ~ Serica spl
Jaynes Tiphia tolopulctala Allen and Chosen China~ IInornaln ieversil
Taynes Phyllopertha pubicolis
Jarabidao A no mala spp CrapedD7ous tillialis sehaum1 Jnpan _~ Serica sppl
Popillia illponica
bull From flelcJ ohsermtion In the second report (10) this species was listed under the Mme of Tiphifl malavalla Sub~quent stUdies
middotof matrilll from Chinn discioSNi the fllct tlllt this WIIS nn undescribed species It lllls since h~en described ilS T fral er (18)
The tachinid Pexomyia aenalis was reared at the Riverton N J laboratory fwm adults of Popillia iaponica collected in central Japan in 1925 and shipped for rem-ings of IIamaria inco7l[Jl1ta Since tbnt time it hus nevel been cncountered in the work in Japnn
Becnuso the work in China wns discontinued at the end of 1926 no further investigations hnve been carried on with Tiphia communis Tfrater 01 the Chinese form of T 7Iotol)olita which so fUl as is known
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
INVESTIGATIONS OF PARASITES OF JAPAXESE BEETLE 29middot
occur only in that country Termination of the work ill India ill 1929 brought to a close all work on T assamensis und T cla1lseni
Rearing work with Tiphia agilis T bicalinata T blelilineafa T koreana T oLidorsalis and T totop1l1lctata wus discontinued beclluseshyof their time of appearance in the field Tlllse species oecm during July and August when the gwbs of Popillia ja]lonica Anomala orientalis A1lioselica castanea and Selica peregrina have pupated or are too small to be nccepted by these parasites
Adult shipments of Oampsomeris annulata were discontinued at theshyend of 1926 Although this scoliid ill oviposit uncI elfmiddotvelop on fullshygrown grubs of Popillia japonica in the laboratory its normal hosts in the field nre of the larger ruteline species pUl-ticularly species of Anomala and Phyllophaga This species hus three genemtions a yenr one of which is present in the field during midsummer when the grubs of P japonica are very scarce in the present nreu of beetle infestation in the Unitecl States
Shipments of the carabid predator Craspedonotus tibialis wereshydiscontinued after 1921 because attempts to establish this speCies w(r( unsuccessful probnbly because of the decided change in the ecological conditions
REARING AND SHIPPING OF SCOLIIDAE
REARIiG SCOLlIDAE
The same general m(thods of rearing scoliid cocoons for shipment were employed as those used in former )poundtus and discussed in previous bulletins (9 10) In India a composition of soil and chopped sphagshynum moss was used in tbe rearing ClOSS sections inst(ud of the usual medium of soil and sod used in Japan nnd Chosen This composition retarded the excessive loss of moisture and improvNl the physical condition of the soil thus increasing cocoon formation A summary of the rearing results obtainNI with the differ(nt sp(cies of Ti[ihia which baw been studied during the period from 1929-32 is giveIl in table 8 It will b( seen from table 9 that of the five speCies of TiTJhia that attack Popillia japonica (T popilliacora T lernaiis T matura T pullicora and Tillhia sp Taiwan 5) a total of 375aS wpre shipped as reared cocoons during the period covered by this bulletin The variance in the ra tio of cocoons prod tlced to the number of ovipositions as between the different sp(cies ranging from 46 percent in Tiphia 7~solata to 800 perc-ent for T saloi may be attributed to differences in temperature find soil conditions pxisting in til( various localities rather than to differences in thp specips themselves
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
30 TEOHNICAL BULLETIN 738 U S DEPT m AGRICULTURE
TABYE S-Rearing results with variolls species of Tiphia
I I i Ratio o( dismiddot 1RntioOfCtlCOOnS i carded comiddot
t I to ovipositions coons to promiddot C t 0middot coCoonj cCooons dutedcocoons
S~cics Of Tiphia fr(u~l~i~h I
0
obtained Ycar i l~ d~~d By c~~~d i B~ By spcdes t B species I I I year Bnd t year nnd 1 1 I sourcr source
----------------1------shy Yllmbtr SlLmlJcriPercrnl Percmt NlLmb Percent Percent
rrhostn 1929 17407 7 (12 4il5 377 53 l do 1930 391)4 15 23~ ~ ( 43 4 3975 I 2Iil 156 do 1931 65hlBmiddot 316ll2 middotI~I 3191 101 J1do i93~ 41549 IS 659 419 i 3793 203middot
T bitrulafa (sccond ~cmrntionloo Japan 001912 38939 S507 218 218 616 bull 721 72
T 1)llrrtlli o-
110 1932 277 61 220 220 I oj 0 1 o T ca3tlttneatrOTa do 19~3 943ti 53~O 571 571 32-1 60 60 T hamonmiddotular do 1932 1101 274 2~9
1
lOi 36 36 T oIaia __ bull do 1932 lfgtlk 76 21~ i ~r 9 ns 11 S T matuTII Todia 192J 24il-lj S012 ~2~ 2 a22 r 212 26 26 T IlOiopolita almi Japan bull 1912 fl3rKmiddot 1594 )middot9 17 0 ~ ZI6 US HS T popilliarora rrho~cn l~aO 2[gt89 474 1 bull ql 1 r 139 29 3
1351 I 259 ~ l 47 i 3 f 102 T bullaloi bull I ~lgmiddot ~l 5 ~m 200 so 0 800 164 520 820 T sfernoia Japun 1932 410 20- 1 4 4~4 ~i 3S 3S
45 7 (qiliptuga lt10 1912 933 3bull1 155 210 lUi 145r mali Chosen 1l~11 65 rr 30-1 504 564 666~ bull 10 ISO 1iphia sp Taiwan l Taiwun ~ li9 911 -12 5 -125 21 213~~ iriphic ~p Taiwan 2 dn 192J 310 277 277 S 93 9 3
Dm-ing- tlI( coursp of rtflring- york with Scoliidnl th( ~-OUll2 lan-ue in tlH ClOSs-sltetion tmys (on1lt in eontnet with various fuctors in till soil which w(akin tlJ(gtm to sueh un (xt(nt that dlllth oftln OC(urs uft(r til( spinuing- of the (o(oons Till mon important ot t1Hse detrill1lntal factors nrl attacks by fungi and disN1S( und ummtisshyfuctorY soil conditions including insufiiri(nt moistun rsuall~- d(ud 111l1t(rial cnll bl distinguish(d byfungus growth and dnrk-11I0wll discolorations on thl (xtrrior of th( (0(0011 Table 8 ulso shows a sumll1l1ry of th( cocoons of the yarious splcils of Tiphia that ([( discard(d at thl tim( oi packing h(cuus( of th( dlnth of th( Inrvne within th( (ocoons
SHIPprVG Cocoo-s -0 AOULT FEMALES
Th( m(thods of handling and ptlcking cocoons for silipm(nt during th( p(riod 1929-33 w(m much til( SI1DW us thos( that hnd gi-(u sutisshyfuctory l(sults in pr(-ious ~middotNllS Extrlml cur( wus tak(11 in s(pnratshylng th( cocoons from the soil in I(uring trays nnd in c-1iminating ull fungus-uff(ct(d disN1Srcl nnd crtlslHd (ocoons prior to pucking
Th( cocoons from Jupan und Chos(n (1( pInc(d -ith n small quantity of slightly moist sphngnum moss in small individual conshytninils 1 inch ill ditlmd(r tlnd 1~ of fin inch dNp Th(s( tins w(re pacl-amiddotd for shipnHnt in u nwdium of slifhtly moist moss in tight ooltl(n CtlS(5 Cocoons (al(d in Tnelia w(n puck(d indiv-idunlly with n ~mfill qunntity of slightly moist sphngnum moss in tin crossshysection sets (Ileh compnrtment of which wns of ubout l cubic ineh
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
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34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
31 ITVESTIGATIOSS OF PARASITES OF TAPASESE DEETLE
Capacity The cross sections were slparated from one another by middotsheets of tin On arrival at the ~roorestown N J laboratory the middotcocoons were tak(n out of their con taitHls plac(d individually in small vials and sto1ld in the temperature-contlol cbambers fOf hibershynation and enwrgence the following s(ason
Yith those species of Scoliidzl( that were suffici(l1tly abundant in the field and that had an aPrtlge longevity of 30 dnys or more it was found to b( more satisfactor~~ und economical to ship adults than cocoons In this cnSl the adult femules are rdease(l in the beetl(shyinfested arens almost imm(diateiy on Rlrh-af whereas with cocoon shipmCnts fiCld rPieus(s cannot b( made until the following season nectssitating the udd(d problPms of overwintering of cocoons and mating
Thp only chunge in thE method of shipping adult f(males from that describ(cl liirliN (10) was in the mattel of food In the early shipshyments from Japan ClIosn and Chinn the food consistcd of a wrnk solution of eitilEr honey or sugar and wutN This was foIlowrd in 1927 nnd 1928 by the usr of a semisolid food consisting of Japn1Hse ngnr mixfd with Cithpr honp~r or sugar syrup in rqual proportions with grratly improHd rrsults During tl1( period 1929-33 still better Irsuts w(re obtained by tlu tIEr of good candy a composition -commonly uSNl as ber food This was made by mixing and knracling powdrred sugar frre of starch into pUfr houry which had bren slightly heated at the rate of 5 pounds of sugar to 2 pounds of hOIHY The Irsulting mixture was packrd into tlir holrs of a woodrl1 food block and wirrd to thr insidr of the shipping Ciln
SrLlARY OF PARASITE srnp~mSTSTO Tn EClTEDSTATES
From tilt incrption of tlle projrct in 1920 to 1933 inclusive 1l1rge numblrs of the Yflrious parasites ill diflrfPnt deveopmrutlll stages have beell shipped to the r nited States In table 9 is given a rr(ord of these shipments The shipments from 1920 to 1928 inclusiv( nrc given as a total for this period while the shipments made during the folshylowing 5-year period are listed separately under each yrar Table 9 shows the country of origin of each parasite und the various stnges of development of the parasites at the time of shipment
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
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CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
fA111]] 9- middotRrcord oj shiplllellts oj jlarusil( lIIaleriallo Ihe Uniled 8laIs for Ihl control oJ tlto J apullcse beett pound11111 relelled 8carall(citiae 192088 CI ~
J tunily genus nnd species 1-3 _____ ~1~~O o~ ~~~(IIIIC~~middot_ _~~J ~__ ~_ i~~~~_=~-~~I~J _~~~J lual 1~21 lIIal J~ tltj (J
Iu(hiultillc I Nt1l1b~r NI~mberjNltI~berNltlllber Nll11lbtr NIpoundIlbc)NuII~r ~ (ollrla dIU lllrllslUled hectles ltIelld bullbull bull I JIIIlIUI 1101 middot1111 I18 (KK) I Uo(XI flIO1 IIIS ~ 1IIrfI)i~ jaeulla rio bull bull ltIn bull SIl bullbull bull bullbullbull SIl
(JIlulIHUirJ liCOIifrWI FI~ltlmiddotcolhmiddotctcd hcclillS liIlug I do IO()middotII~) I j bullbull j -- aOO4()O[rolhols dOIlell F-PUIHlrI bullbull i do bullbull middot1middot j 217 I bull 217 tlyr~olidue lduMiIu J~rlu do lillll 17102 I~OO L 1 21402 td
Dcxildllll qlJrriu Iollull lnmsltiled grubs Chosou bullbull middot1 1)middot middot1 r)()() 2OroO 4000 I~_ [c) 12-1
JIIIlIIU iliiCXl mOIJ() 000 315001ro1 slIirUu ~ielilcOIIcted gruls bull do IflUOOO 150000 ~
~((lIidlle Adult lomnlos J JlIllIn Chosen Chinn 310110 alOliO ~
(I(lmlomtri~ ulluuluiu_ Hmlred cucoons ~~ _ ~w _~ _ bullbull 348 ~ doIIIrnsltl1Od ~rubs bull do 01n 1 1 6410 Tida uUI~ nHrlld C(WOOUS ~ IChOeIL~~~~~ _ 1U7 1 107
Adult l(lIllllie 9004 10511 1I2nO CO rilhifl ufericur 00 l~
-
Hcorcd CU(OOIlS_ 02 lOS Tiptiu bkurillttla I do bullbullbullbullbull ryen~~~~i5~I2i8 2S4Il148~0 57O
Adult ICIllIlIcs bullbull I1l51 aonTiphu 1Ii~Ilutt ~ Hellred cocoons J3()J 7692 bullbullbullbull 20725 [Iphlu bredilleaa UI ll Uliiphlll bllrulli 01I ~ middotmiddot1middot~~t~)middotbull~rphia Cn~tlJlItlIClora lt10 bullbull 5066
Adult lemllies bull J071671~ -middotmiddotIlt~I 5~~1JiphifJ rommlLs ~ HOllred cocoons j do 1480 I 1180
Altlult lemllics bullbull do 1216liphill Ia bullbull J Hellreltl cocoons bull do 2l0 ~ iphu hO(ll(lIlW(lri 1 do 2(14middot I Jnplln I~~middotImiddot~middot ~~~~middoth~~ ~~~ ~~ oiJihill Isolalll do 711 njJlphiu 1IH1lurn 10 middot I~~ 2i4~ 7800 I middotmiddot1middotmiddotmiddotmiddot 352iO
Pillhiu WloJJoillJ do 11 liphiIlJlpolilI ulflli do bull ~~j JUpllll Chosen_ ~ I rmiddotjmiddotIiiImiddot Iaili ()JJhu oliuorHJlif lt10 bull t Chason 119 0 ifhill lhglolhIlYllc do I Chosen Chinn c bull 14011
AltlUlt lemnlllS _ j JflPan Chosen Chinn ~Jm 1gt1 middot1 rmiddot 10 niphlll Jvplllliltora
Htured tocnons ~ _ bull do 10 ii4 bull bull bullbull I 35 I 644 I 1 ai~a3 r1 riJlin pulliora 1 VhmiddotIltImiddot(olieclltlltl cocoons 81Oi7 tllndill 811157 bull I I [phibullbullaol bullbull 11AtlUIt IcnllllllS ChoJ~n I bull bullbull bull bull 4(1 I middotHI 1-3
bull i Henrad cocoons r1 i Altlult IcmlllllS Jnpnn~_ 4$il i1 I 44171JiJhiu fallulu 8U bull
~~ tltjbullbull middot1 Henred cocoons bull lt10 ~~JUI bullbull 1middot 200 Imiddot middotmiddot
7ipllia tuilploga I lt10 do ) I 1180 11801iphill O(I)III(IIa do Chusen bull ~~ _ ~~ ~ I 476 ___ _ I ___ ~~laquoj~_w ___ ~ ~_~ ~~_ _~ ___ middot170
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
1iphia (mlliis bullbullbullbull_bullbullbullbullbullbullbull_bullbull Adult (cnllllo~ --- bullbullbullbullbullbullbullbull bullbullbull -bullbullbullbullbullbullbullbullbullUeurcd cOloons bullbullbullbullbull _bullbullbullbullbullbullbullbullbullbullbull
[phi( sp 1lIlwDn 5do bull bullbullbullbullbullbullbullbull bullbullbull riphiu p lolwnu 2_ ~ __ __ ~M _ w bullbull __ tlo ~~ ~ ~ __
Cllmhltlnc CrapltdOllotIUJ libanbullbull bullbullbull Adult heeties bullbullbullbull _bullbull
------_
5718 I 11120 I 12412 60211CI)o~~)n~~~~IlI~ 1 ~ 1i~ 1~~ 085j ~~~o 283amp1 498li 1li1U_ _ ~~+~ ~_~ ___ r~_~4_ ain ~ ___ _ 371i
bullbullbullbull lt10 bullbullbullbullbullbullbullbullbullbullbullbullbullbullbullbull bullbullbullbullbullbull 39 bullbullbullbullbull au JIl[Jnn 10450 16450 -------_-_ bull--------------shy__
~ 3 fJ
S ~ o ~ en o ~
~ en
~ en o I-zj
~ ~ ~
~ gj ~
~ ~
CIl CIl
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
34 TECHX[CAL BULLETI~ i3S U S DEPT m AGRIClTLTURE
SUMMARY
This bulletin is hfiSPcl on the rpsults of further iuVCstigations OIl
the natunll enemips of Popillia japonica and rdated Scalllbneidae in Jnpnn Chosen India and Taiwtln (Formosa) during the ppriod from 1929 to 193~ inrlusin-
Innstigations Wlle carried on with four sppcies of Tnchinidae find one species of PTrgotida( parnstitic on adult Popillia l)(etles Ilnd with two species of DexiidnC and fin splcies of 8coliidtle pnmsitic on the grubs of Popillia
IJamalia incolItlla Illld Trophops ria1l8eni hnH- two or more generashytions it year in JiLpan and an prc(nt ill the fidd as adult onr n longel pCriocl than til( Popillia Iw(tiPs find so nquire altNnllte hosts
elllet( tinEN( the mot lfrCctiC piuusitl of Popillia aponica in northlII1 Japnn was found to be well distributed on the i5lnnd of Hokkllido when it (ffects a high mtpound of parnsitizntion (ach YNllmiddot On til( island of Honshu just south of Hoklmido it was found most abundnnt in the hill and mountninous sections of tht nolthelll and Wlstlrn portions of the island nlthouglt its prespnce in ynrious locnlishyti(s was ditinctly nriabk and the degllp of parasitization wns conshysidpmbly 10w(I tlltln tlltlt found in Hokkaido C ciner((pound WitS ulso found in th( mountainous section of Kyurhu the most southe1l1 of the larglI islttnd of In pnn From 1929 to ] 933 a totnl of 33663 pupitriu Wlf( rearNI nnd hipp(d to thp l~nit(d SlatCs
Ilamaria IlC( ngnlfl which OlCUlS ~Pilelally throl1hout th( Oripnt WtlS found in mun n(w loculi tils on the islunds of Honshu Shikoku and Kyushll in Japan but thp glntlIll rate of fipld parnsitization wit exreectingly low and yuriaole
No fUItlwr l(Cords of ElIiJmiddotopsi~ jamna as a pnrnsitp of Popillia were obtained Trophops claweni wns found in senral n(w localitils in western Japun but the percentnge of pnlllsitizution ill P jap()ni((~ wns eIT low
Fllrtilpr r(co[(ls of Adapsiliajladstla n pyrgotid pnrnsite of Pop ill ia beptles in India rCY(al(d the same low rat of ptlrnsitization in lhe field as bad been preiously reportNI In 1929 it total of 3800 pllparia of this parnsitp weI( reared lind shipped
Considemblp naring work was ClllTiNl 011 with thp bo clpxiicis DeJmiddotia zmiddotpntrali8 in Cllosen and Prouna sibirilfl in Japan During this period 48900 Pnpiltia gruhs W(Ie pnrnsitized in the lnborntory by thrs( 2 dpxiicl and shipPNI to till lnited Stategt
Imestigotions were carriNL on with flY( specils of Tiphia parnsitic on Popilli(pound jILponim T pnpillialol(L nnd T lfrnalis occur in both Jtlpall and Chospn while T malum anrl T pllllil)f(L occur only in India Thl prps(ncl or thps sppcips in nhundance in the field was always localized somrtin1(s chan~ing from nnr to YPUImiddot bpng sermin~lY (kpPIHI(nt on a suffirilnt supply of adult food tlnd suitabl(middot host glllhs Extensin rlnring work wns Cllrrird on with thts( spprils of Tiphia and a total of n)88 1parpd cocoons welp shipPNI during the 5-ylttH plIiod In ndclition 47Sl) adult f(mllips of T lfrTlaliR
were colJrctrd in tlH ipld tlnd shipp((1 of which npPlOximntc-ly 84 percent arrived in tIll CnitNI Statl ole
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
f
t j
_ - shy__ middot_rmiddot
is
j
L
r
L
INVESTIGATIONS OF PARASITES OF JAPANESE BEETLE a
During the cOUlse of th( investigations 011 the pamsites of Anomala orientalis AutoseriNL castanea und Serica peregrina no suitabl( parnsit(s of tIl( adult bl(tl(s w(le disco-(red A large number of sppcles of Tiphia pnmsitic on grubs of Anomala nnd Serica W(re found but onl 11 sp(cips were consicllN[ of sllffieirnt importance to WaTllnt introduction Of thes( Tiphia bi8eClllata T bllrrflli and T (middotgilishyplaga occur only in Japun tlnd are parnsitic on Anomala whil( T noto]J()lita alieni u1so a pnrmi t( of Anomala occurs also in Chos(n tlnrl China T ca~tallea(rora T i~olata tlncl T 8tel1w[([ nl pnmit(s of Saica occur only in Japan hil( T Clspricae and T liomoncula ri8 u1so parasitp of Sericrz occur in both Jltpatl and ChosPl1 T satoi which is -rlY dosp to T sternaa tnxonomically is 0 pnmsitr of S(Iica nnd occurs onh- in Chosen Of all tll(gt Tipltia stlldiN[ in Jnpltll and CbOSP11 only T biReclilata and T tegitiplaga haY( two g(nprations a y(ar Lik( th Tiphia pnrnsitls of PIJpillia th(se splcis of TiphilL ar ypry Iocalizrd nnd tlllil ((f(ctinns and distriblltion al( d(pltndshyent on a suIficiPllt snpply of adult food and suittlblc host glubs Shipmrnts of adults and rentcel cocoons of these 11 sprcies flom 1929 to ]933 incIusin totaled 613911dult females and 77270 cocoons
LITERATrRE CITED (1) ALLEX H und JyXES n A
1 028 PROGRE~ WITFI TilE DIPORlED PARiiITES OF THE IAPAXESE BEETLE DlmiddotPIXC 192 Tour rCOll Ellt 21 823-832 ilhlH
(2) --_ and JAYXE H A 1030 COXTnmrTIO 10 THE TAXOXO~IY OF ASIATIC PS OF THE rEloTS
rIPHIA (SCOl[[DAF L S Xatl ifu1 Pmc 76 art Ii 105 pp ilIll
(3) --- and n[RRELL R Y I 1134 )IETHOOi OF OBTAIXIX E~IER(aX(E OF TUlilA ADrLTR FWHI ntshy
PORTfD (OC00X FOR OlE AGAINST TfiE IPAESE BEETLE Jour Ag-r R( 49 000-922 iIIus
(I) RUOCK J V- 1931 THE TApe OF TIPlllA VERAUf ItOHWfm AN 1~IPORTED 1RRITE
OF THE IAPgtESE BEETLE AT THE CI01E Ot 1913 Jour Ecoll Flit 27 middotHll-496 illu~
(5) nRDRO -L H ID34 Tin FLCTtATIOX OF TilE pOPtLAT10X OF TIp1IIA POpILLlVORA
ROHWEll IX THE FIELD AXD IT POSSIRLE CAFSE-i JOur Ecoll Ent 27 )1+-518
(6) BCRRELL R Y 1931 DEX VENTRALIS AlDRHmiddotU AN DIPORTED lAnMHTE OF THE JPAXESE
BEETLE Jour Agr Hcs 43 323-336 illlli (7) (A~IERO P
1910 DECRlpTIOSS OF 3 XE FOfOHIA[ HnIEXOPTEHA FRO~I RORXEO Ent nund~chau 28 12-130
(8) (1AoSEX C P 1931 IXSECTR JXJrRIOCS TO AGRICeLTenE IN JApAX C So Dept Agr
(ir 168 116 pp ilus (9) --- 1IXG r h and TERlmMIT (
1927 THE PARASITES OF POPILLlA IAPONICA IN JUAX AXD CflOSE (KORE) AXD THEIR INPRODrCTIOX INTO THE CNITED STATES r S Dlpt A~r Bul 1middot129 56 pp iIIUll
(10) --- JAYXES H A and GARDXER T RoO 1933 FlCRTIlEIi INVESTIGATIONS OF THE PARASlTS OF POPILLIA JPOXIC-
IX THE FAR EAST r S Dept Agr Tech BIl 366 59 ppill us
36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
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36 TECHXICAL nULLETIS i 38 U S DEPT OF AGRIClLT1RE
(11) GARDNER T R 1934 (OlllARArlE onpOSITIOX EnI(IENlJY Axn (OLIE(TlOX (OSTS OF
DIIORJo~D ER~eS ESTABLISHED lllH[A VERXI[S HOl[WEII A pHMilTE 01 rUE JAIAXERE BEEIIE Jollr Econ Ellt 27 middot197-199
(12) [IOLWIL Y J K 1931 lEliPERATURE AS A A(lOR 1- ~IIE ACTIVITY AND DEYELOP~[EXI
OF THE CIllNESE STIUIN O~ 11lIllA IOlllLIA YOIU (nOI1W) IX XEW JEREY -LD pENNSYLYAINA X Y Ent SocJour 39 355-565 illus
(13) K[N J 1 1931 THE pREEXT SlAl( OF THE ESIAurISIIED PARASITE 01 pOP[LLlA
lAIOX[( NEIl IIX Jollr Eroll Ent 24 middot153-middot162 illus (14) --- ALLEN H W Ilnd HALLOCK H C
1927 IHE PHESExr STNITS OF TIH WORK 00 filE PARASITE OF POPIlLIA JAIONLCA XEID[AX Tour Ecoll Ellt 20 a65-373
(15) --- and 11-LLOCK HAROLD C 192)t HEpOR1 00 IJoHrAlX pAItASI1ES OF pOP[[IIA JApON[CA NEWlI bull
Jour Eco Ent 18 30L-3j6 illml (16) --- and HOLLOWAY bull1 K
1930 THE ElBLISIDIEN1 AND (OLONIZAl[ON OF TIII1[A POPILLIAIOH A PAR-tSIrE 01 TilE HI XESE HEE1LE Jour Eron Ellt 2a 26poundi-27middot1 illu-
(17) --- and HOLLOWA Y T K 1930 TIPIUt POpILL OIU ROHWER A 11 RSITE OF THE IAIAo ESE llEETLE
L R Dept Agr Cir 1-13 12 pp illus (18) PAHKER L B
1935 TIlHEE XEW SPE(IE 01 [IIH 11l0l[ EASTEUX AlIA X Y Ent ~oe Jour 43 39[middot -lOol
(19) 1937 SEEX NEIl RIE(IE M ASIATIC T[PIIIA X Y Ent Soc Jour
45 [2j9j-290 (20) HOHWEH S A
1924 DECIUPTIOX OF TIIREE NEW SPECIER OF TIFlilA IAIUlITIC O~ IOPILLIA JPONI(A (lID) Bnt Soc Wash Proc 26 87-92
(21) ROBERTS HAHlo-on 1 930 SEVE~ XEW NAl[Ei l~ THE Gms rIIIIIA (RYlIEXOPTERA SCOLIIshy
DAE) Canad Ent 62 189-190
-------~-----~---- - bull - ~--- ------~ +~- --~---~-~
01 snlgt by tll~ fIP(middotjnPIHl~nt of Oocllllcnr~ iURhillgron O Cbullbullbullbull Price 10 cents
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