16 Dr. Hugh Scott on Some Nycteribiidae from the

SOME NYCTERIBIIDAE FROM THE AUSTRALIAN REGION PART I : SPECIES FROM THE NEW HEBRIDES

By HUGH SCOTT, Sc.D., F.L.S., F.E.S.

THIS and the following paper include descriptions of two species of bat-para- sites new to science and a record of a third species from the Ken- Hebrides, an archipelago whence no Nycteribiid had previously been made known ; a description of a new species from Fiji ; and remarks supplementing earlier descriptions or bearing on the synonymy and geographical distribution of several other species from the Papuan region, New Caledonia and Australia. These last are mainly species of Cyclopedia, and i t is not attempted to give an account of the entire Nycteribiid fauna of the countries under review, except in the case of the New Hebrides.

I have examined material from the New Hebrides collected in the islands of Espiritu Santo and Tanna in 1985 by Dr. P. A. Buxton, in Espiritu Santo in 1927 by Dr. J. R. Baker, and in Malekula in 1929 by Miss L. E. Cheesman. Three species are represented, namely, (i) Penicillidia buztoni, sp. n., allied to P. ocmnica (Bigot), a form occurring in New Caledonia and Queensland, the host of both these species being Miniopterus australis ; (ii) Nycteribia (Listropodia) bakeri, sp. n. (also from Miniopterus austrazis), intermediate between two species, one of which is known from the Moluccas, Amboina and parts of Australia, and the other from New Caledonia, the Loyalty IS. and Queensland ; and (iii) Cyclopodia ozycephala (Bigot), previously known from New Caledonia and the Loyalty Is., now recorded for the first time from the New Hebrides.

The new species from Fiji, Cyctopodia pewzbertolzi, is allied to C. oxycephala, the distribution of which has just been mentioned, and to C. minor, Speiser, which is known only from New Britain. As to the other species discussed, Penicillidia oceanica was previously known only from New Caledonia, but is now recorded from Queensland. The identity of Nycteribia ptwopus, Rain- bow (1904), with Cyclopodia albertisi, Rondani (1878), is established, and New Guinea, Salwati and the Kei Is. are added to the known range of this species, previously recorded from the Moluccas and parts of Australia. C. sirnilis, Speiser, is very close to C. albertisi, but its distinguishing features are given after examination of part of the original type-material, and the Timorlaut Is. are added to its range. The original description of Cyclopodia minor, Speiser, is supplemented. I am indebted to the Museums of Berlin and Genoa for the loan or exchange of material which has helped to elucidate the relationships of this confusing group of species of Cyclopedia. Several of the new records are based on material sent me many years ago by the late Dr. Frederick Muir, others on specimens more recently received from various sources by the British Museum.

The eight species reviewed in these two papers are all, so far as known, confined t o the Australian zoogeographical region, but, if Australia proper, the Austro-Malayan (Papuan) and the Polynesian be accepted as subdivisions of that region, most of the species occur in more than one subdivision. The Australian region as 2 whole seems relatively rich in species of these parasites, though it is difficult to estimate how i t will eventually compare with the Orlental and Ethiopian regions."

* Musgrave (1925, Rec. Austr. Mus., 14 : 289-300, and 1927, 15 : 263-276) has enumerated seven species recorded only from Australia proper ; eight species are revieis ed

Australian Region. Part I : Species f r o m the Xeio Hebrides. 17

Returning to the critical study of NPCTERIBIIDAE after an interval of about five pears, I am more than ever impressed by their bizarre forms and structural modifications, which place them among the most remarkable of all insects. The varying degrees of reduction of segmentation in the abdomen of the female, the extraordinary shapes assumed in some cases by the chitinous sclerites, and the variety of combinations of these shapes presented by different species, are indeed striking. There are also groups of species in which successive gradations in the development or reduction of particular characters can be clearly traced. Instances of these phenomena are mentioned in the present paper under the heading ( ( Affinities ” in Penicillidia bzixtoni and, in the follow- ing paper, in the remarks on Cyclopodia sitnilis. Individual variation is discussed in the following paper under C’yclopodia albertisi.

The iyplpes of the new species are in the British Museum. The material examined is all preserved in alcohol, and the drawings have

all been made from specimens lying in alcohol, with the help of a drau-ing- apparatus. Figure 1 is from a drawing made some years ago in Cambridge by Mr. H. Gillings, the remainder are from detailed pencil-drawings by myself, inked over and in part completed by Miss 0. F. Tassart. I am much indebted to these two helpers for their care.

PENICILLIDIA, Kolenati. Penicillidia buxtoni, sp. n.

Resembles P. oceanica, Bigot, in several respects, notably in the extreme reduction of the abdominal ctenidium, but differs from that species by its larger size, also (in the d) by the form of the claspers, and, in the 9, by the completely different arrangement of the abdominal segmentation, especially on the dorsal surface.

Head not laterally compressed, bare a t the back and sides, but with the vertex bearing moderately long bristles, behind which are short bristles, extending further back than in P. oceanica. Eyes, a single minute ocellus on either side, just below the bristly area, absolutely the same colour as the surrounding chitin, and only visible when the head is viewed in profile under a high power. Thorax with the ventral plate appearing about 15 (in the d), nearly 1; (in the q), times as broad as long, convex from front to hind margin; median longi- tudinal suture impressed and dark-coloured, curved transverse sutures inconspicuous, surface covered with very fine short bristles, hind margin with about eight bristles on either side of the middle, of lengths varying from very short to moderate. Dorsally, the lateral ctenidia appear normal, but are largely hidden (when viewed from above) by a fold of the whitish connexivum (as in P. oceanica and other species of this genus), only the distal pert of the front six or seven teeth being visible ; there is an oblique transrersc series Of about seven rather long bristles on the chitinous part of the thorax, on either side, just in front of the halter and level with the front of the hind coxa. Legs rather long and slender, the pseudo-articulation near the base of the femur, and the bristling,

in the present paper and its sequel, and eight others have been recorded from various parts of the region, so that twenty-three is the total so far known for the Australian region. For comparison it may be noted that I enumerated seventeen species from India, Ceylon and Burma (1925, Rec. Ind . Nus., 27 : 351-384). Professor G. F. Ferris recently expressed verbally the view that the Austro- and Indo-Nalayan faunistic areas are the part of the world richest in KYCTERIBIIDAE, possibly their centre of distribution, and that the poverty of Korth and South America is real, not merely due to American forms being still undis- covered.

Length (front of thoracic sternal plate to end of abdomen) m. 2.5 mm.

18 Dr. Hugh Scott on Sowe Xycteribiirlne front the

being as indicated in the figure. .ibdomen. j : True basal tergite small, bare, not clearly separated from second; the folloir ing tergites are densely bristled ; surfaces of tergites 2-5 thickly covered with fine, short, erect bristles, the bristly area on the fifth being very short ; tergites 3-6 have their chitinous parts progressively shorter. and in the sixth this part is so short that there is no room for surface-bristles except a t the sides; marginal bristles moderately long on tergite 2, from moderately to very long (and strong) on tergites 3-6; anal segment bare abore in its basal half, but with erect bristles of moderate length on the posterior half of its dorsal surface and on the sides, long outstanding bristles on either side before the hind angles, and a series of fairly long bristles across the hind margin. Ventrally, the ctenidium along the margin of the basal sternite is reduced to about twelve

1 mm.

FIQ. 1 .-Penicillidia buxtmi, $, dorsal view.

very short and rather fine teeth, widcIy spaced, and alternating nearly regularly ni th ordinary bristles, which are moderately long in the middle and very long towards the lateral margins; surface of basal sternite bare right a t the base, otherwise rather closely covered with fine short bristles, decumbent and directed backwards except a t the sides, where they are erect and turned outnards. Sternites 2 and 3 have the marginal series rather irregular, two, three or more short bristles stand between each two long ones, and the surface is bare in the middle except for a sparse irregular series of fine short bristles in front of the marginal series; a t the sides the surface is covered with short erect bristles and the margins bear very long bristles, some erect, others directed backwards. Sternite 4 has an irregular transverse group of short, sharp, teeth on the middle part of its hind margin, the median basal part of the surface is bare, and the hind part bears rather sparse,

Australian Region. Part I : STecies from the Xeiu Hebrides. 19

short, erect bristles with very long ones a t the sides near the hind n i q i n . Chspcrs with their outer margins sinuate, not regularly arcuate as in P. oceanica (cf. fig. li in the description of that species by Falcoz, cited below).

FIG. L.-Penicillidia buxtoni, 6, ventral view of thorax and abdomen (scale 2 larger than fig. 1).

FIG. 3.-PeniciZZidia buxtoni, 9, dorsal view of abdomen (same scale as fig. 2).

9 : Basal trrgite rather sniall and narrow, widening from its base to about the middle of its length, then tapering to an obtuse apical angle which bears from two to four rather long bristles, apart from which it is entirely bare; it nearly divides into two halves the

20 Dr. Hugh Scott on Some Nycteribiidae from the

chitinous part of the large second tergite, the surface of which on either side is densely covered with short erect bristles, while the hind margin has a series of from eight to ten moderately long bristles on either side and a group of about seven in the middle. Behind this is a large area of whitish connexivuni, bare except a t the sides, where it is fairly densely covered with bristles of moderate length ; it bears two nearly quadrangular chitinous plates, each with about six long stout bristles on its hind margin, the inner ones very long; a few short bristles stand just in front of the marginal series, the rest of the surface of each plate is bare ; behind these a transverse median fold is visible in the connexivuni ; anal segment short, tapering, subtruncate, with surface above mostly bare, erect bristles of medium length a t base, on its sides, and across the hind margin, those a t the hind angles being very long; the anus itself is on a very short narrower extension, projecting behind the marginal series of bristles as a chitinised ring fringed with fine hairs. Ventrally, the basal sternite has its ctenidium even more reduced than in the $, there being only a few very short teeth on either side and none in the middle, where two or more short, fine,

FIQ,. 4-Penicillidia buxtoni, 9, ventral view of abdomen (same scale as fig. 2).

ordinary bristles are set between each two long ones; surface bare a t the base, otherwise rather sparsely covered with short fine bristles, all decumbent and directed backwards ; behind this is a large area of connexivum, bare at the base, then densely covered with short bristles and with a curved transverse series of eight very long bristles. Behind this, two nearly ovoi’d chitinous plates, their hind margins set with bristles of varying lengths, their surfaces bearing six or seven erect bristles (on each plate) just before the hind margin, the outer ones of which are very long. Behind these, a short transverse chitinous part,* with hind margin closely set with moderately long bristles, short erect bristles on the surface just before the margin, and long (or very long) erect bristles in a dense group on either side. Subgenital plate bilobed, densely covered with erect bristles, those a t the apices of the lobes being very Iong.

N E W HEBRIDES : Espiritu Santo, Hog Harbour, 21.vii.1925, from Mini- opterus australis, Tomes (Oldfield Thomas det.), 1 8 and 1 $? (types) and 6 9 paratypes (one bearing an almost completely extruded larva) ( P . A . Buzton) ;

* The outline of this transverse chitinous area is not very sharply defined and is not indicated in fig. 4.

Ausirnlinn Regiot?. Part I : Species f r o m the New HebTides. 21

Tnnna, S3.is. 1915, from i7liwiop/~~ixs ait.s~ralis, 1 2, 2 4 (P. A . Ric&o~) ; Espiritu Santo, Hog Harbour, li.ii.1927, from Minioyierus alts/rcclis, 2 $, S C ( J . R. Bnker).

This species and P. oceanica constitute an extreme develop- ment along a certain line within the genus Penicillidia. In the great reduction of the abdominal ctenidium they approach the genus Bre),loctenia, Scott, the type (and hitherto only known) specics of which, E. pryressa (Muir), Scott, occurs in Amboinn on iVi?ziop/~ri/s.* P. buxfoni and P. oce(mica, however, retain some vestige of the abdoniinal ctenidium, while they have the lateral thoracic ctenidia normally developed ; but these latter are partly hidden under folds of the connexivum, which may possibly be a step in the direction of their disappearance (they are quite absent in Eremoctenia). The presence of ocelli (organs entirely wanting in Eremoctenia) also helps to determine the position of these two species in the genus Penicillidia.

A more elementary stage in the reduction of the abdominal ctenidium is seen in the widespread Oriental species, Penicillidia jenynsi , Westwood, the teeth being in that species rather far apart in the middle. P. jenynsi and P. bzixtoni also exhibit some resemblances in the form of the abdomen in both sexes, notably in the underside of the $? abdomen; cf. Scott, 1908, Trans. Ent. SOC. Lond., 1908 : 360-362, pl. xviii, figs. 1-5.

i4finities.

Penicill idia oceanica (Bigot). Nycleribin ocennica, Bigot, 1883, Ann. Sac. ent. Fr., 1885 : 21G. Penicillidia oceanica (Bigot), Falcoz, 1023, in Sarasin et Roux, Xovn Cdklonin (ZOO].), 3, (i) :

86-89, figs. 5-8.

Distribution : New Caledonia ; Queensland. The original description, based on specimens from New Caledonia, without

record of host, was inadequate, and the species was fully redescribed and figured in both sexes by Falcoz, again from New Caledonian material, some of the specimens having been found on Jliniopterus australis, others on an undetermined bat ; I had previously compared examples with Bigot’s original types for Dr. Palcoz. Two records from Queensland can now be added: Mossman, from “ illiniopteriis schreibersi,” t 1 $, 2 $? (Frederick Muir, NO. 471) and Townsville, 1927, 1 $, 2 9, without record of host (G. Dennes).

These are the first authentic records of the species from Australia, since the record of a single $ from New South Wales, published by Speiser in 1901, has been shown by Falcoz not to refer to this species.

NPCTERIBIA, Latreille. Subgenus LISTROPODIA, Kolenati. Nycteribia (Listropodia) bakeri, sp. n.

Very close to, and in some respects intermediate between, N . (L.) parilis, Walker (Scott, 1914, Ann. Mag. Nat. Hist., (8), 14 : 230-234, pl. xii, figs. 20-24), which is known from Aniboina, the Moluccas and Australia, and N . (L.) sarasini, Falcoz, 1923 (Nora Cale‘donia (Zool.), 3, (i) : 89-93, figs. 9-12), which is recorded from New Caledonia, the Loyalty Is. and Queensland.

Length (front of sternal plate of thorax to hind end of abdomen) approximately 1.75 mni. Thorax with sternal plate about as broad in its widest part as i t is long; its hind margin bears some longer bristles, including one very long bristle on either side just within

* 1917, Parasitology, 9 : 600-GO5, pl. xsiv. 7 This almost certainly refers to Niniopferus uztslrulis, no\\ treated as a distinct species.

22 Dr. Hugh Scott on Some Nycteribiidae from the

the hind angle (as in N . (L.) parilis). Legs : the form (typical of the subgenus) and pro- portions of length to breadth are shown in fig. 5. Abdomen. The diagnostic characters are indicated in fig. 6 and can be best stated in tabular form, which will also serve to show the differences between the three species.

N . parilis. N . bakeri. AT. surasini. Both sexes.

First sternite with pos- First sternite with pos- First sternite with pos- terior row of surface bristles terior row of surface bristles including about four to six fairly short, of equal length. which project beyond the apices of the ctenidial teeth.

terior row of surface bristles fairly short, of equal length.

Claspers longer, more slender, arcuate towards apex, which is acute.

Surface of 3rd sternite with relatively few bristles, that of 4th sternite almost bare.

Tergite 2 with surface al- most bare, marginal bristles weaker.

Membranous part of ter- gite 3 fairly widely covered with scattered moderately long bristles.

Anal segment with two separate dorsal chitinous plates, their surfaces bare, a curved series of short thorn- bristles along the hind mar- gin of each (the two series convergent in front).

0" only. Claspers shorter and

broader, nearly straight but convergent, with inner mar- gin obliquely cut away to- wards apex, which is sub- truncate.

Underside very bristly, the surfaces of sternites 2, 3 and 4 all covered with numerous short erect or sub-erect bristles. (The very long erect bristles on the hind margins, four on sternites 2 and 3, two on sternite 4, are very marked.)

? only. Tergite 2 with surface-

bristles fairly close in mid- dle part, marginal bristles stronger and longer.

Membranous part of ter- gite 3 fairly widely covered with scattered moderately long and strong bristles ; one or two postero-lateral bristles on either side situated apart from the rest, on the mem- brane.

Anal segment with two separate dorsal chitinous plates, each with two trans- verse series of 2-4 (usually 3) short thorn-bristles in front of the middIe.*

Claspers shorter and broader, convergent, with apical portion bent up- wards and apex emarginate, almost bifid.

Underside less bristly, surface of sternite 4 almost bare.

Tergite 2 with surface glabrous, marginal bristles relatively short.

Membranous part of ter- gite 3 with only a small part of the surface bristled.

Anal segment with two separate dorsal chitinous plates, the short thorn- bristles forming a single irregular transverse series and apparently variable in number.

* I n one example there is only one transverse series.

Australiait Region. Part I : Specks from the New Hebrirltx. 23

The most definite characters of hT. balceri lie in the very distinct forin of the 2 claspers, and the long bristles on the hind part of the surface of the basal sternite, which project beyond the teeth of the ctenidium in both sexes. In the 9, the arrangement of the thorn- bristles on the dorsal chitinous plates of the anal segment is much closer to that in N . sara- sini, but the bristling of the middle part of the surface of tergite 2, and of the membranous portion of tergite 3, more nearly resembIes the arrangement in N . pasilis. The two very long marginal bristles near the hind angles of the sternal thoracic plate are of about the same length as those of N . parilis, and longer than those of N . sarasiiii.

1 mm. t I

FIG. B.-Nycteribia (Listropodia) bakeri ; from top to bottom, right front, middle and hind legs, viewed from in front.

NEW HEBRIDES : Tanna, 22.ix.1925, from Miniopterus australis, Tomes (Oldfield Thomas det.), 2 3 and 3 (inchding the 8 and $2 types) (P. A . Buzton) ; Espiritu Santo, Hog Harbour, 153.1927, from Miniopterus atutralis, 3 2 and 4 ?, also 1 8 from the same place and the same species of bat, 2O.iii.1927 ( J . R . Baker).

CYCLOPODIA, Kolenati. Subgenus CYCLOPODIA, s. str., Scott, 1917, Parasitology, 9 : 607.

Cyclopodia oqcephala (Bigot).

Ngcteribia osycephala, Bigot, 1860, Ann. SOC. ent. Fr. (3) 8 : 227. Cyclopodia oxycephakz (Bigot), Speiser, 1902, Z. Hymenopt. Dipt. 2 : 170; Falcoz, 1923, in

Sarasin et Roux, Nova Calbdonia (Zool.), 3, (i) : 03-96, figs. 13-16; id., 1924, Bull. Rlus. Hist. nat., Paris, 30 : 19.

Distribution .- Loyalty Is. (Lifu) ; New Caledonia ; New Hebrides. Thc host of the New Caledonian material is recorded as Pteropus ornatus, Gray.

Originally inadequately described by Bigot, this species was placed in its right generic position by Speiser, and has been fully re-described, with figures of the dorsal and ventral surfaces of the abdomen in both sexes, by Falcoz.

It has not previously been recorded from the New Hebrides, but the follow- ing records from that archipelago can now be published : Espiritu Santo, Hog

t I

i mrn

FIG. 6.-Nyderibia (Listropodia) bakeri ; from left to right, ventral view of 6 abdomen (with apex of one clasper more highly magnified), dorsal view of 9 abdomen, ventral view of $2 thorax and abdomen.

Harbour, 17.i.1927, on " N'Kara Kara " (Pterops eotinus, I(. And.), 1 $, 1 ( J . R. Baker); Elephant Island (a small islet off the coast of Espiritu Santo, near Hog Harbour), 1927, from three examples of Pteropus geddiei, MaoG., respectively 2 $, 1 9, and 1 $, 1 9 ( J . R. Baker); Malekula, Ounua, iv-v, 1929, from fruit-bat (species undetermined), 2 3,1 $?(Miss L. E. Cheesman).