Transactions of the Royal Society cfS. Aust. (1993), U7(2>, 97-104. FEMALES AND 1MMATCJRES OF THE AUSTRALIAN HYALOPSYCHE DISJVNCTA NEBOISS (TR1CHOPTERA), AND A NEW FAMILY PLACEMENT

by A. WELLS* & D. CAKTWRIGirrt

Summary

Welis, A & CurrwKicwr, D. (1993) Females and immature* of the Australian caddisfty Hyalopsyche (Usjuncm

Neboi8* fTrichopterai. and a new family placement, Trans. /?. Sac. S. Aast. 117(2), 97-104, 4 June. 1993. The female, larva and pupa or the northeastern Australian Hyalopsyche dixjuncta Neboist;, 1980, an: figured and described for the first time. Comparisons between Hyabspxyehe disjwuui and several North American species

of' PIvAocentropus Banks. 1907 (Family Dipwudojuidae Ulmer, 1904), show that both genera share apornorphouK lar»al features. Transferral of Hyalopsyche Ulmer, 1904, from the family Hyolopsychidae Lestage, 1925, to the fondly and supression of Hyalopsychidac are proposed.

Key Words: Tricnoptera> Dipseudopsidae, Hyalopsyche, larvae, pupae, females.

Introduction numerous adults of both sexesw and just prior to departure, made (he exciting find of several larvae and The history of unstable of the group of pupae Female and larva] and pupal features of this polycentropodid-like caddisfly genera including species are here described and illustrated, and the larval Dipseudopsis Walker, 1832, Pratodipseudcpsis U liner, niche is described. For comparisons, some larval 1904, Banks, 1907, and Hyalopsyche feaiuies of Phylucemwptis srx, and the female genitalia Ulmer, 1904, is discussed in detail by Weaver & and head showing the tentorium of the North American MaJicky (in ms). These authors present strong support P. iucidus (Hagen) axe also figured. for the familial status of Dipseudopsidae, including the Confirmation of sharing of the specialised larval first three of the above-mentioned genera „ and possibly niche and other synapomorphous larval character also Hyalopsyche, an African-Oriental genus, which states, firmly supports the monophyly of placed by Schmid (1980) with Phylocentropus in MM Phyloeentrvpus — Hyalopsyche, and we present an the family Hyalopsychidae. Convincing evident* for hypothesis for tbe derivation of die specialised female the monophyly of Dipseudopsidae Ulmer, 1904, sensu genitalia of Hyalopsyche from those of Phylocentropus. Weaver & Malicfcy. derives from a consideration of We argue that Hyalopsyche should be placed in the features of the female abdomens and larvae. family Dipseudopsidae sensa Weaver & Malieky (in Although reference has been made in the past to ms), and the name Hyalopsychidae is here supressed, females of Hyalopsyche (Ulmer (1915) identified three Ross (1965) and Ross & Gibb* (1973) discussed the New Guinean female specimens as H, rivalis

Northern Territory Museum of Arts and Sciences, PO Box Phylocentropus sp were provided on loan by Dr J. 4646, Darwin, N.T: 0801. Present address: Australian Morse. Biologic*! Resources Srudy, GPO B°x WG> Cafthetra, ACT Specimens were prepared for study by treuimen* in 2601. Biology laboratory, Wem'bee Treatment Complex, Private caustic potash, or using a bcto-phenol preparation. Bag ta Werribee,' Victoria 3030, Drawings were made using camera lucidas on a Wild 98 A. WELLS & D. CARTWRIGHT

Figs 1-3. Hyalopsyche disjuncta Neboiss, female: I, dorsal head showing tentorium; 2, mesothorax, dorsal view; 3, genitalia, lateral view. Figs 4-6. Phylocentwpus lucidus (Hagen), female; 4, dorsal head showing tentorium; 5, mesolhorax, dorsal view; 6, genitalia, lateral view. Scale bars = 1 mm, unless otherwise indicated. .

FEMALRS ANO iMMATCRBS OF THE AUSTRALIAN CADDISFLY 9*

M5 dissecting microscope and a Wild M20 compound brown, dorsal margin of prothorax and strengthening microscope. ridges of thoracic pleunj black* rest of body pale. Body Material b deposited in the following institutions: length 60-14.8 mm. head capsule width (160-088 mm. Australian National Collection. Canberra Head round in dorsal view (Fig. 12); fronro-clypeus (AN1C); Museum of Victoria, Melbourne, (IWV); sub-rriangular, extending to posterior of head capsule, Northern Territory Museum of Ans arjd Sciences, anterior margin rounded, tentorial pits situated Darwin, (NTM); and Queensland Museum, Brubane, posteriorly; labium broadly rounded with few setae, (QM). Anterior margin finely divided. Mandibles Cartwoght and A_ head capsule, palpi absent. Prothnrax only WellS NMV; 2 acr. 1 9. IT 39'S 142° 28'E, with sclerotiscd tcrgjte. Fore trochantin fused to pleuroa (Fig. IS). Mesosternum McDonnell-Cockatoo Creek. jn T l«.ii.l992> D, Cartwright and A. Ufells, NTM; D cr o\ 5 9 0, 31° (Fig. %) with a pair at elongate sclerites posteriorly.

44 S 142° 29 'E> Gunshot Creek, at Telegraph Legs (Figs 1&. 19) short flattened , tarai with dense Ciosving, I7.ii.l°92. Q Cartwrighl and A. Wells, QM. fringes of setae, claw on fore limb strongly curved. Fore tarsus ith adpressed row 11 Of cr, 8 9 9- same loc. and collectors, t4-tS.iU992. M an of modified setae QM; 3 crcr, 15 9 9, same loc and collectors* on the mesial margin. IIU9.ii.1992, NMV; 21 o'er. 4 9 9, *ame loc.. Abdomen elongate, with a lateral fringe of fine setae, 4-5.io.1992, M. Grassland, ANIC: 31 cror, 19 9 9, wiihout grlis. but with a group of anal papillae (Fig. anal prolegs elongate, with of long same loc and collector. 10-U.iv.i992, NMV; I pupa, 7); cluster setae same loc, D. Cartwright and A Wells, !8.iv 1992, anterior to the slender strongly curved anal claw (Fig. NTM; 2 larvae, same loc and collectors, NTM; 7 20). larvae U reared to aduk 9 >. 1 pupa, same Joe. and Pupa (Figs 9-lL 21). Length 7.5-1(13 mm Labrurn collectors, 19.ii.1992. CFig. 9) short, with a small marginal papilla apJco- Neboiss (1980) illustrates the general body features tnesally. Mandibles (Fig. 10), simple, slender, curved plates (Fig. 3-5 of male Hyatopvjche dvijancm , and gives anterior wing Hook 21) with denticles, on abdominal length as 6 mm, but in our samples male anterior wing segments Hl-Vin. posterior places on segment V only, length ranges from 4.3-6.2 mm. Females conform with developed into hand-like processes, fbsterior abdomen males in general features, including reduced maxillary (Fig, 10 divided into two simple setose lobes. palpi and warts on the lucsothofa* (Fig. 2). bat are Biology. Larvae and pupae were extracted from considerably larger. An addition;*] feature, the fragmented silken tubes. The exact architecture of the tentorium, is figured (Fig. l>: its anus arc well larval dwelling is unknown, but the branched portions developed. v>llh inner lateral flanges posteriorly, and of tubes appear to have all the features of the tubes lhe posterior bridge complete of Phyhcentropus piacidus (see Wallace el at 1976) ftmale (Figs 1-3). Actetroi wuig length 6.0-8JQmm. and P. carohnus Carpenter (see Wiggins 1977, Fig, Genitalia in the tbrm of a slender, elongate, noa- 15 4D). (Such features are also shared by retntctjlc ovipositor (Fig. 3). Sternne VIU not Pnotodipsettdopsis sp. (un-named) as illustrated hy subdivided. external gonopod ot segmenl VJJ1 Uender, Gibbo (1968, Fig, 3)). The architecture of the tapered, reaching almost In tip of segment EX. Hyafopsyrhe disjuncta dwelling is probably ainulaj ft&teriorly segment X is divided m for m a pair of distal a blunt-ended vertical branch with a small opening lobes, each with apical cercus and one papilUi p^TlinHy projected into the flow, a series of connected cross fused on inner margin, the second projecting veoually tunnels, and a dilated net chamber. from the outer margin at about the level of division Habitat. Larval tubes were collected front a small of segment X. stream with a rather unstable base of coarse sand . The

Larva (mature* (Figs 7, 8. 12. 13, 16\ 38-20|. dwelling tunes were found attached to logs amongst

Cainpodeiforui, slender, elongate, (Fig. 7). Head, deeply packed sand aod litter, usually in situations pronotum, limbs and anal claws brown-yellow or pale where the litter, trapped against logs embedded in the 100 A. WELLS & D. CAKTWRIGHT

Figs 7-8. Hyalopsyche disjuncta Neboiss, larva: 7, whole , lateral view; 8, mesosternum.

Figs 9-1L Hyalopsyche disjuncta Neboiss, pupa: 9y labrum; 10, mandibles; H, posterior abdomen, Scale bars = 1 mm, unless otherwise indicated. FEMALES AND 1MMATURES OF THE AUSTRALIAN CADDISFLY m

13

Figs 12, 13. Hyalopsyche disjuncta Neboiss, larval head, dorsal and ventral views. Figs 14, 15. Phyhcentropws sp., larval head, dorsal and ventral views. Scale bars = 0.1 mm. 102 A. WELLS & D. CARTWRIGHT

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Figs 16, 17. Hyatopsyche disjuncta Neboiss and Phylocentropus sp. , larval mandibles,, dorsal view. Figs 18-20. Hyalopsyche disjuncta Neboiss, larva. 18, foreleg and proleuron; 19, midleg; 20, anal proleg. Fig. 21. Hyalopsyche disjuncta Neboiss, pupal hook plates, right side. Scale bars =0.1 mm. FEMALES AND IM MATURES OF THE AUSTRALIAN CADD1SFI.V m

sand, was tegitining in decompose. Collections nf in the wing*; as well as other more major ones, such adults were obtained at lights beside small and larger as shape of head, position of antennae, reduction of streams; the smaller streams generally were almost maxillary palpi and form of genitalia These differences completely shaded by the canopies of riparian

maxillary palp or H. disjtwcui is broad and apioally McLachlan during this study, indicated that cm the basis truncate rather than tapered and rounded; labia are of female genitalia and wing form, Pirytocensroptts and modified to form virtually identical slender, elongate Hyalopsycht are closer than Dipscadapsis- spinnerets Thoracic features, too, show close Hyoicpsyche. resemblance and the highly specialised legs differ only Association or females and larvae of a species in in the exact shape of the specialised setae adpresscd Hyaiopsydxe has helped to stabilise the higher level to the niesal margin of the fore tarsi. H. disftwaa lacks taxonomy of this genus, placing rt in the femUy gills but conforms with Pkyfocesttropus species in fbrtn Dipseudopstdae. On the basis of studies reported heTe of anal ptolegs and their claws. The larval niches and examination of the literature, we suggest that appear to be identical. Hyaiopsyche and Phylocewropas represent sisier Pupae are relatively uninfomiative: mandibles of groups, H. frawNifti resemble those illustrated by Ross (1944, Ross &. Gibbs (1973) suggested on the basu crt Fig, 204) for R placidta. Little information is available similarities between Phylo-crumpm and on pupae of Rkyloccntropux. Polycentropodinae that Phyloccmropus is closer id the Some of the above features occur in other caddis ancestral stock than other Dipseudopsinae larvae. R>r example, specialised elongate spinnerets (=Dipseudopsidae| and, given its occurrence in the are seen in larvae of some P&yebomyiidae and some Oligoccne Baltic ambers, they suggested t Eurasian Bcuouudae, although in these families the spinnerets origin tor tne subfamily (=famUy). We have not found are proportionally shorter than in Rkyloct'nJropas and evidence with which to dispute this contention — the Hvalopsyche. Simple ana] claws *Jso occur elsewhere, single north-eastern Australian species in Hyalopsyefie but are probably plesiomorphous. The highly modified, is undoubtedly of Oriental origin. Careful phylogeoetic flattened limbs, the form oi the mandibles and the larval analyses in the future will elucidate intrafamilial niche are (bund only in //yafryifycAe, PfjylocertJrofMis. relationships, but *l last the family Dtpseudopsidae is Pwtodfpseudopsis and Dipseudopsis. reunited. The probability of men a suite nf resemblances Aclitinwledgmcnts evolving by convergence are slim. The simplest hypothesis is thai these tasa arose from die same W& thank the Royal Geographical Society of specialised ancestral stock. Queensland for providing the opportunity and financial Adult characters do not concur with those of the support for field work at °Heathlands** and the larvae. No clear synaponiorphies can be recognised, surrounding Jardinc River 3rca of Cape YotK, and for which is lite x

r' has been confused for so long. Among adult features members of the Society during our stay at *Heathlands several small differences in general body features can D, Carrwrigbfs work was undertaken with backing be observed between Hyakptyche and PHytocentropus, from his family, and A. Wells with backing from the such as the arrangement of setae on the meta-*cutellum Northern Territory Museum of Arts and Sciences, (sec Figs 2. 5) and die absence of several crossvetns Darwin. We are grateful to both parties. 104 A. WELLS & D. CARTWRIGHT

Adults of and Dr J,S. Wfeaver of the University of New Hampshire, comparisons. USA, kindly provided material of Phylocentropus larvae of Phylocentropus sp. were made available by University, USA. lucidus and Dipseudopsis indica from his collection for Dr J. Morse, on loan from Clemson

References

Dipseudopsinae (Trichoptera, ). /. Banks, N. (1907) Descriptions of new Trichoptera. Proa the Soc. 8, 312-16. Ent. Sot: mshington 8, 117-133. Georgia Em. Insectes et Arachnides du Canada, Betten, C. (1909) Notes on the Trichoptera in the collection Schmid, F. (1980) "Les 296 of the Indian Museum. Rec. Indian Mus. 3, 231-242. partie T. (Agriculture Canada). pp. Westafrikanische Trichopteren. Gibbs, D.G. (1968) The larva, dwelling-lube and feeding of Ulmer, G. (1904) Uber 353-359. a species of Protodipseudop.sis (Trichoptera: ZooL Anz. 28, besonders von Ceylon Dipseudopsidae). Proc. Ent. Soc, London 43, 73-79. (1915) Trichopteren des Ostens, Dtsck. Entomol. Z., 1915, 41-75. Lestage, J.A. (1925) Notes trichopterologiques. VIII. Etude und Neu Guinea. Catalogue of the specimens of du groupe polycentropidien et catalogue systematique des Walker, F. (1852) in the collections of the British genres et especes decrits depuis 1907 dans Genera neuropterous 65: 93-120. Museum. Pt. 1, 192 pp. (London). insectorum. Bull. Ann. Soc. Entomol. Belg. } W.R., Staats, A.A. (1976) Neboiss, A. (1980) First record of the subfamily Wallace, J.B., Woodall, & Hyalopsychinae from Australia (Trichoptera: The larval dwelling-tube, capture net and food of ft>Iycentropodidae). Fblycentropodidae). Arch, fur Hydrobiotogie 90, 357-361. Phylocemropus placidus (Trichoptera: 149-54. Ross, H.H. (1965) The evolutionary history of Ann. Entomol. Soc. America 69, (In ms) The genus Phylocentropus (Trichoptera: Psychomyiidae). J. Kansas Weaver, J.S., HI., & Malicky, H. Asia (Trichoptera: Dipseudopsidae). Em. Soc. 38, 398-400. Dipseudopsis from "Larvae of the North American (1944) The Caddis Flies, or Trichoptera, of Illinois. Wiggins, G.B. (1977) (Trichoptera)". (University of Toronto BulL Illinois Nat. Hist. Survey 23, 1-326. Caddisfly Genera & Gibbs, D.G. (1973) The subfamily relationships of Press: Toronto and Buffalo).