Journal of lnsect Behavior, Vol. 5, No. 6, 1992 Short Communication Transfer of a Larval Mantispid During Copulation of Its Spider Host Sonja J. Scheffer t' z Accepted May 18, 1992; revised May 18, 1992 KEY WORDS: Climaciella brunnea; rnantispids; Schizocosa species behavior; spider parasites. INTRODUCTION Within the Mantispidae (Neuroptera), larvae in the subfamily Mantispinae feed on the eggs of spiders. Two methods are known by which larval mantispids gain access to spider eggs. In some species, the larva actively searches out a spider egg sac, penetrates the silk covering, and enters the sac. In other species, the larva climbs aboard a living spider and remains there until an egg sac is produced. During the construction of the egg sac the larva leaves the spider and enters the sac. Particular mantispid species may be obligate egg sac penetrators, obligate spider-boarders, or facultative penetrator/boarders, using either strategy (Redborg and MacLeod, 1983). While mantispid larvae aboard spiders are known to feed on the hemolymph of their hosts, development beyond the first instar does not proceed until feeding begins on spider eggs (Redborg and MacLeod, 1984). Because only female spiders construct egg sacs, natural selection in spider- boarding speci es should favor larvae that preferentially end up on female spiders. Selective boarding of female spiders has not been detected in the mantispid species studied so far (Redborg and MacLeod, 1985; S. Scheffer, personal observation). A possible alternative to selective boarding of female spiders is the transfer by mantispid larvae from male to female spiders during copulation. Transfer behavior during copulation was not found in Mantispa uhleri Banks (Redborg and MacLeod, 1985), and it has not been looked for in any other species. ~Department of Biological Science, University of Cincinnati, Cincinnati, Ohio 45221. 2Current address: Department of Ecology and Evolution, State University of New York, Stony Brook, New York 11794. 7 9 7
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8/4/2019 Transfer of a larval Mantispid during copulation of its spider host
Journ al o f lnsect Behavior , Vo l . 5 , No. 6 , 1992
Short Communication
T r a n s f e r o f a L a r v a l M a n t i s p i d D u r i n g C o p u l a t i o no f I ts S p i d e r H o s t
Sonja J . Schef fer t'z
Acce pted M ay 18 , 1992; rev ised M ay 18 , 1992
K E Y W O R D S : Climaciella brunnea; r n a n t i s p i d s ; Schizocosa s p e c i e s b e h a v i o r ; s p i d e r p a r a s i t e s .
I N T R O D U C T I O N
W ith in the M ant i sp idae (N europ te ra ), l a rvae i n t he sub fami ly Mant i sp inae feed
on the eggs o f sp ide rs . Tw o me thods a re know n by w hich l a rva l man t i sp ids
ga in access to sp ider eggs . In some spec ies , the la rva ac t ive ly searches out a
spider egg sac , p ene t ra tes the s i lk cover ing, a nd ente rs the sac . In o ther spec ies ,
the la rva c l imbs aboard a l iv ing spider and remains there unt i l an egg sac i s
p roduced . D ur ing the cons t ruc t ion o f the egg sac the l a rva l e aves the sp ide r and
ente rs the sac . Par t icula r mant i sp id spec ies m ay be ob l iga te egg sac pene t ra tors ,
obl iga te sp ider-boarders , or facul ta t ive pene t ra tor /bo arders , us ing e i ther s tra tegy
(Redborg and M acL eod , 1983) . W hi le m ant i sp id la rvae aboard spiders are kno wn
to feed on the hemolymph o f t he i r hos t s , deve lopment beyond the f i r s t i n s t a r
does no t p roceed un t i l f e ed ing beg ins on sp ide r eggs (Redborg and MacLeod ,
1984).
Becau se only fem ale spiders co ns t ruc t egg sacs , na tura l se lec t ion in sp ider-
boa rd ing spec ie s shou ld favor l a rvae t ha t p re fe ren t ia l ly end up on fem a le sp ide rs .
Se l ec t ive boa rd ing o f f ema le sp ide rs ha s no t been de t ec t ed i n t he man t i sp id
spec ies s tudied so fa r (Redborg and MacLeod, 1985; S . Scheffe r , pe rsona l
observa t ion) . A poss ib le a l te rna t ive to se lec t ive boarding of female spiders i s
t he t r ans fe r by man t i sp id l arvae f rom ma le t o fem a le sp ide rs du r ing copu la t ion .
Trans fe r behav io r du r ing copu la t i on w as no t found in Mantispa uhleri B a n k s
(Redborg and MacLeod , 1985) , and i t ha s no t been looked fo r i n any o the r
spec ies .
~ D e p a r t m e n t o f B i o l o g i c al S c i e n c e , U n i v e r s it y o f C i n c i n n a t i , C i n c i n n a t i , O h i o 4 5 2 2 1 .
2 C u r r e n t a d d r e ss : D e p a r t m e n t o f E c o l o g y a n d E v o l u t i o n , S t a te U n i v e r s i ty o f N e w Y o r k , S t o n y
The pur pose o f th i s s tudy wa s to inve s t iga te the poss ib i l i t y o f t r a ns f e r
be ha v io r in the m a n t i sp id C l i ma c i e l l a b r u n n e a ( S a y) , a n ob l iga te sp ide r - boa r de r.
M E T H O D S
C . b r u n n e a , a m i m i c o f P o l i s t e s wasps (Wheele r , 1889; Ople r , 1981) , i s
f ound f r om the we s te r n a nd c e n t r a l Un i t e d S ta t e s sou th to C e n t r a l Am e r ic a
(Opler , 1981) . C . b r u n n e a l a r va e on m a le sp ide r s we r e ob ta ine d f o r th i s s tudy
i n tw o w a y s . I n t h e s u m m e r o f 1 9 8 7 , a n u m b e r o f a du l t m a n ti s pi d s e m e r g e d
f r o m t h e e g g s a c s o f l a b o r a t o r y - m a t e d f e m a l e S c h i z o c o s a o c r e a t a ( He n tz )
( Lyc o s ida e ) , wh ic h ha d b e e n c o l l e c t e d a s suba du l ts a t t he C inc inna ti N a tu r e
C e n t e r ( H a m i l t o n C o u n t y , O h i o ) . T h e s e m a n t i sp i d s w e r e a l l o w e d t o m a t e ; s u b -se que n t ly , e ggs we r e l a id on the w a l l o f a p l e x ig la s s c on ta ine r by a s ing le f e m a le .
F o l lowing ha tc h ing , t he m a n t i sp id l a r va e we r e a l lowe d to boa r d S . ocrea ta
sp ide r lings , wh ic h we r e the n r e a r e d in the l a bor a to r y un t il t he f o l lowing sp r ing
wh e n the y r e a c he d m a tu r i ty . Add i t iona l m a le sp ide r s (S . ocreata a nd S . rovner i
Ue tz a nd Donda le ) c a r r y ing m a n t i sp ids we r e c o l l e c t e d in M a y o f 1988 ( a t t he
C inc inna t i Na tu r e C e n te r a nd in no r the r n Ke n tuc ky , r e spe c t ive ly ) . The m a n t i s -
p ids , o f t e n pos it ione d on the sp ide r ' s pe d ic e l o r c a ra pa c e ( R e dbor g a nd M a c Le o d ,
1983 ; S . S c he f f e r , pe r sona l obse r va t ion ) , a r e r e a d i ly e v ide n t . A l l m a n t i sp id -
be a r ing sp ide r s u se d in th i s e xpe r im e n t c a r de d on ly one m a n t i sp id .Tw e n ty - th r e e pa i rs o f m a le a nd f e m a le sp ide rs w e r e p la c e d ind iv idua l ly in
15-ml p las t ic cul ture d ishes and le f t for 3 to 4 h unt i l copula t ion or , in two
c a se s , un t i l c a nn iba l i sm oc c ur r e d . O ne m e m be r o f e a c h pa i r c a r d e d a m a n t i sp id ,
wh i l e the o the r d id no t . F o l lowing c opu la t ion a l l sp ide r s we r e c he c ke d f o r the
p r e se nc e o f the m a n t i sp id . I n no c a se wa s m or e tha n one m a n t i sp id f ound pe r
pa i r .
R E S U L T S
F our te e n c opu la t ions oc c ur r e d in wh ic h the m a le wa s in i t i a l ly c a r r y ing a
m a n t i sp id . I n 12 in s t a nc e s ( 85 .7%) the m a n t i sp id wa s f ound on the f e m a le a nd
wa s m is s ing f r om the m a le f o l lowing c opu la t ion . I n 2 o f the se 12 c a se s the
f e m a le a t e the m a te be f o r e I ha d de te r m ine d w he the r o r no t the l a r va ha d t r ans -
f e r r e d du r ing c opu la t ion . Whe n the se f e m a le s ha d f in i she d f e e d ing the y we r e
c he c ke d f o r the p r e se nc e o f the l a rva . I n bo th c a se s the m a n t i sp ids ha d t r a ns-
f e r r e d suc c e ss f u l ly . B e c a use l a r va e a r e known to t r a ns f e r f r om a c a nn iba l i z e d
sp ide r to i ts p r e da to r [ two a dd i t iona l c a se s in th i s s tudy ; R e db or g a nd M a c L e od
(1985) wi th M . u h l e r i ] , i t i s unc le a r w he the r the se two t r a ns fe r s oc c ur r e d du r ingc opu la t ion o r c a nn iba l i sm . Af te r d i sc a r d ing the se two c a se s , 10 suc c e ss f u l t ra ns -
f e rs r e m a in o f 12 c opu la tions ( 83 .3 % t r a ns f e r) . La r va l t r a ns f e r wa s ne ve r se e n
wh e n the in i ti a l c a r d e r wa s a f e m a le sp ide r (n = 9 ) .
8/4/2019 Transfer of a larval Mantispid during copulation of its spider host
L a r v a l M a n t i s p id T r a n s f e r B e h a v i o r 7 9 9
L a r v a l t r a n s f e r b e h a v i o r w a s d i r e c t l y o b s e r v e d i n o n l y o n e c a s e . D u r i n g
c our t sh ip , a nd a t t he s t a r t o f c opu la t ion , t he m a n t i sp id r e m a ine d f a i rly s t a tiona r y
on the pe d ic e l a nd po s te r io r po r t ion o f the c a r a pa c e ; th i s i s t he usua l c ond i t ionf o r m a n t i sp ids on sp ide rs no t e nga ge d in m a t ing - r e la t e d a c t iv it i e s. D ur ing c op -
u la t ion , t he m a n t i sp id be c a m e ve r y a c t ive , r e pe a te d ly m ov ing r a p id ly a r ound
a nd und e r ne a th the m a le c a r a pa c e un t i l i t f i na lly d i sa ppe a r e d unde r the m a le ; i t
wa s l a t e r f ound on the f e m a le .
D I S C U S S I O N
B e c a use l a r va l m a n t i sp ids se e m no t to boa r d f e m a le sp ide rs p r e f e r e n t i a l ly ,
a g r e a t num be r o f l a r va e m us t boa r d m a le sp ide r s , t he r e by l a c k ing d i r e c t ac c e ss
to e ggs ( w h ic h a r e r e qu i r e d f o r de ve lop m e n t be yon d the f ir st i n s ta r ). Th e t r a ns f e r
o f l a r va l m a n t i sp ids f r om the i r m a le sp ide r hos t s to f e m a le sp ide r s du r ing c op-
u la t ion i s a n im por ta n t s t r a t e gy , a l lowing the l a r va e to pos i t ion the m se lve s to
e n te r e gg sa c s.
Th i s i s the f i r st r e por t o f l a rva l t r a ns f e r be ha v io r du r ing hos t c o pu la t ion in
a m a n t i sp id spe c ie s. T he t r a ns f e r be ha v io r shown by C. brunnea wa s no t de te c te d
in a p r e v ious s tudy invo lv ing M . u h l e r i ( R e d b o r g a n d M a c L e o d , 1 9 8 5 ) .
T h e d i ff e re n c e in t r a n s f e r b e h a v i o r b e t w e e n C . b ru n n e a a nd M . u h l e r i c ou ldb e d u e t o a n u m b e r o f f a c t o r s . M . u h l e r i i s a facul ta t ive sp ider -boarder ; i t i s
a b le t o e n t e r p r e v i o u s l y co n s t r u c te d e g g s a c s . R e d b o r g a n d M a c L e o d ( 1 9 85 )
sugge s t t ha t, f o l lowing the de a th o f it s m a le sp ide r hos t , a l a r va m a y be f r e e to
wa nde r in se a r c h o f a n e gg sa c ( o r a ne w hos t ) . Th i s a l t e rna t ive r ou te p r ov id ing
a c c e ss to e ggs i s no t ope n to C . b ru n n e a , w h i c h d o n o t r e c o g n i z e p r e v i o u s ly
f o r m e d e gg sa c s o r e ggs a s f ood ( R e dbor g a nd M a c Le od , 1983) . S e le c t ion f o r
l a r va l t r a ns f e r be h a v io r m a y be m or e in t e nse in a spe c ie s in wh ic h suc c e ss f u l
l a r va l de ve lopm e n t a bso lu te ly r e qu i r es tha t a l a rva i s c a r ri e d by a m a tu r e f e m a le
spider .An o the r f a c to r tha t m a y in f lue nc e the e vo lu t ion o f l a rva l t r a ns f e r be h a v io r
i s the num be r o f sp ide r hos t s u se d by e a c h m a n t i sp id spe c ie s . The hos t r a nge
o f M . u h l e r i i s ve r y b r oa d ; i t ha s be e n c o l l e c t e d f r om sp ide r s in a t l e a s t e igh t
f a m i l i e s ( R e dbor g a nd M a c Le od , 1985 , Ta b le 27 ) . I n c on t r a s t , C . b ru n n e a h as
be e n c o l l e c t e d e xc lus ive ly f r om sp ide r spe c ie s w i th in the f a m i ly Lyc os ida e
( G e o r g e a n d G e o r g e , 1 9 7 5; R e d b o r g an d M a c L e o d , 1 9 8 3; L a S a l le , 1 9 8 6; th i s
s tudy) . R e s t r i c t ion o f C . b ru n n e a t o sp ide r hos t s w i th in a s ing le f a m i ly m a y
pr ov ide a c ons i s t e n t c ue ind ic a t ing hos t c opu la t ion tha t i s no t a va i l a b le to M .
uhler i , whic h use s a f a r m or e d ive r se g r oup o f sp ide r s a s hos t s . Add i t iona l ly ,
l y c o s i d c o p u l a t i o n i s c o m m o n l y o f a l o n g d u r a t i o n , a l l o w i n g a m p l e t i m e f o r
t r a ns fe r . R e s t r i c t ion o f C . b ru n n e a t o th i s s ing le sp ide r f a m i ly m a y ha ve m a de
the e vo lu t ion o f l a r va l t r a ns f e r be ha v io r m or e l ike ly .
8/4/2019 Transfer of a larval Mantispid during copulation of its spider host
T h a n k s a r e d u e t o C o l e G i l b e r t f o r i n i t ia l l y i d e n t i f y i n g th e m a n t i s p i d s .
G e o r g e U e t z k i n d l y p r o v i d e d l a b o r a to r y sp a c e . D o u g F u t u y m a , L e o S h a p i r o ,
a n d K e i th K a r o l y m a d e v a l u a b l e c o m m e n t s o n t he m a n u s c r i p t . A S i g m a X i
G r a n t - i n - A i d o f R e s e a r c h s u p p o r te d m y r e s e a rc h o n r e p r o d u c t iv e b e h a v i o r o f
Schizocosa s p id e r s f ro m w h i c h t h is p ro j e c t sp r a n g . T h e C i n c i n n a t i N a t u r e C e n t e r
a l l o w e d m e t o c o l l e c t sp i d e r s f ro m t h e i r g r o u n d s .
R E F E R E N C E S
George, L. D ., an d George, N . L. (19 75 ). Notes on the three hundred and fifty-eighth meeting: A
new record of mantispid reared from spider. Pan-Pacif Entomol. 51: 90.LaSal le, M. W . (1986 ) . Note on the mant ispid Climaciella brunnea (Neuroptera: M antispidae) in
a coastal marsh habitat. Entomol. News 97: 7-10.Opler, P. A. (1981). Polymorphic mimicry of polist ine wasps by a neotropical neuropteran. Bio-
tropica 13: 165-176.Redborg, K. E. , and MacLeod, E. G. (19 83) . Climaciella brunnea (Neuroptera: Man tispidae): A
mantispid that obligately boards spiders. J. Nat. Hist. 17: 63-73.Redbo rg, K. E., and M acLeod, E. G. (1984). M aintenance feeding of first instar man tispid larvae
(Neuroptera, Man tispidae) on spider (Arachnida, Araneae) hemolymp h. J. Arachnol. 11: 337-341.
Redborg, K. E . , an d MacLeod, E. G . (1985) . The developmental ecology of Mantispa uhleri Banks(Neuroptera: Mantispidae). III. Biol. Monogr. 53: 1-130.
Wheeler , W. M. (188 9) . Two cases of insect mimicry. Proc. Wisc. Nat. Hist. Soc., April, pp.217-221.
