Transactionsofthe Royal SocietycfS. Aust. (1993), U7(2>, 97-104. FEMALES AND 1MMATCJRES OF THE AUSTRALIAN CADDISFLY HYALOPSYCHE DISJVNCTA NEBOISS (TR1CHOPTERA), AND A NEW FAMILY PLACEMENT by A. WELLS* & D. CAKTWRIGirrt Summary Welis, A &CurrwKicwr, D. (1993) Femalesand immature*ofthe Australian caddisfty Hyalopsyche(Usjuncm Neboi8* fTrichopterai. and a new family placement, Trans. /?. Sac. S. Aast. 117(2), 97-104, 4 June. 1993. The female,larvaand pupaorthenortheastern AustralianHyalopsychedixjuncta Neboist;, 1980, an: figured anddescribedforthe firsttime. ComparisonsbetweenHyabspxyehedisjwuuiandseveralNorthAmericanspecies of'PIvAocentropusBanks. 1907(Family Dipwudojuidae Ulmer, 1904), showthatboth generashareapornorphouK lar»al features. TransferralofHyalopsyche Ulmer, 1904, from the family HyolopsychidaeLestage, 1925, to the fondly Dipseudopsidae 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(heexciting findofseveral larvaeand The history ofunstable taxonomy ofthe group of pupae Female and larva] and pupal features of this polycentropodid-like caddisfly genera including speciesareheredescribedandillustrated,andthelarval DipseudopsisWalker, 1832,Pratodipseudcpsis Uliner, niche is described. For comparisons, some larval 1904, Phylocentropus Banks, 1907, and Hyalopsych&e feaiuiesofPhylucemwptissrx, andthefemalegenitalia Ulmer, 1904, is discussed in detail by Weaver andheadshowingthetentoriumoftheNorth American MaJicky(in ms). Theseauthorspresentstrongsupport P. iucidus (Hagen) axe also figured. forthefamilial statusofDipseudopsidae, includingthe Confirmation of sharing of the specialised larval firstthreeoftheabove-mentionedgenera„ andpossibly niche and other synapomorphous larval character MalMsopHlyaacleodpsbyychSec,hmaindA(f1r9i8c0a)n-wOirtihenPthayllogceennutsr,opwuhsicihn Pshtaytleose,entfrivrpmulsy—sHuypaploorptssychteh,eanmdowneopphryesleynt aonf the family Hyalopsychidae. Convincing evident* for hypothesisfortbederivationofdiespecialisedfemale themonophylyofDipseudopsidaeUlmer, 1904, sensu Weaver & Malicfcy. derives from a consideration of gWeenitaarlgiaueoftHhaytalHoypaslyocphseyfcrhoemsthhoousledofbePhyplloacceendtrionputsh.e features ofthe female abdomens and larvae. family Dipseudopsidaesensa Weaver & Malieky (in Although reference has been made in the past to ms), andthenameHyalopsychidaeis here supressed, femalesofHyalopsyche(Ulmer(1915) identified three Ross (1965) and Ross & Gibb*(1973) discussed the NewGuineanfemalespecimensas H, rivalis<Be*teu, phylogenetic relationships of Dipseudopsinae (in the 1909)). no descriptions offemales are available, and Psychomyiidae and subsequently to the until now no larvae have been associated. Rolycentropodidae), but with more data available, A single Australian species of Hyalopsyche, ft phylogeneticanalysesofhighertaxaofTrichopteranow disjunda Neboiss, was described by Nebnws (1980) an progress (Weaver & Frazer pers. coram.) sliould from two adult males from northern Queensland, provide a more definitive assessment of the Repeatedeffortstocollectmorespecimens, including relationship* of thi* family. larvae, failed; H. disjunda appeared to be rare. Recently, however, duringan intensivecollectingtrip Materials and Methods oftwoweek'sduration in the.fardine River areaoftar northern CapeYork (aspart oftheRoyal Geographic Adult Hyalopsychedkjuncw were taken mainly at Society of Queensland's uWet Season" Cape York I2vor 8v blackorUV lights; adults and larvaewere PeninsulaScientific Expedition), dieauthors collected collected intoand stored in70% ethanol. Comparative materialoftheNorth AmericanPhylocentropustucidus (adults) was provided by Dr J.S. Weaver, and adults ofP.pktcidusandOipseudopsisindieusandlarvaeof NorthernTerritoryMuseumofArtsandSciences, POBox Phylocentropus sp were provided on loan by DrJ. 4646, Darwin, N.T: 0801. Present address: Australian Morse. Biologic*! ResourcesSrudy,GPOB°xWG>Cafthetra, ACT Specimens werepreparedforstudy by treuimen* in 2601. Biology laboratory,Wem'beeTreatmentComplex, Private caustic potash, or using a bcto-phenol preparation. Bag ta Werribee,' Victoria 3030, Drawingswere madeusingcamera lucidas on aWild 98 A. WELLS & D. CARTWRIGHT Figs 1-3. HyalopsychedisjunctaNeboiss, female: I,dorsalheadshowingtentorium;2, mesothorax,dorsalview;3,genitalia, lateral view. Figs4-6.Phylocentwpuslucidus(Hagen),female; 4,dorsalheadshowingtentorium; 5, mesolhorax,dorsalview;6,genitalia, lateral view. Scale bars = 1 mm, unless otherwise indicated. . FEMALRS ANO iMMATCRBS OF THE AUSTRALIAN CADDISFLY 9* M5dissectingmicroscopeandaWild M20compound brown, dorsal margin ofprothoraxandstrengthening microscope. ridgesofthoracicpleunjblack* restofbodypale. Body Material b deposited in the following institutions: length60-14.8mm. headcapsulewidth(160-088 mm. Australian National Insect Collection. Canberra Head round indorsal view (Fig. 12); fronro-clypeus (AN1C); Museum of Victoria, Melbourne, (IWV); sub-rriangular, extendingtoposteriorofheadcapsule, 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 <Frg. 16) s:out. dissimilar, each with differing numbers of Systematica denticlesapicallyandon innerdistal margin,andwith adense pcruciUus medially on inner margin, antero- fyttiapwhf ditjuncta Ncboiss, 1980 lateralmargiascrenulate; righi mandiblerugoseantero- FIGS 1-3, 7-13, 16, 18 21 dorsally. On ventral head (Fig. 13), anterior ventral HHoylaolioyppseycnhweleS.sjNunaQaueeNnebsoliasnsd,, W2H9iOv*.l93?597,-36N1M;VF.igs 1-8. aspuobtwoemnciu(mgulrao)unsdhoerdt,anftoerrimoirnlgy,apnaiirseodscseelteasetsriitaunagtleed; Materialexamined:N. Queensland: I Cf Ua 50'S subapically; maxillary palps relatively slender, lobus 142° 30'E. Bertie Creek, at Telegraph tCrossing, iiwernusbroad,obliquelytnmcate indorsal view, and 5.li.l992t D. Cartwrigbi<mdA- Wells, NTM; 2 V OP, bearinga brush ofhair apically: labium modified to 2 9 9, 11° 50'S W2° 30'E. Dulhunty River, «l form a slender, elongate spinneret, about *A as long Telegraph.Crossing,8-9.ti.1992*I> CartwoghtandA_ as head capsule, labial palpi absent. WellS NMV; 2 acr. 1 9. IT 39'S 142° 28'E, Prothnrax only with sclerotiscd tcrgjte. Fore McDonnell-Cockatoo Creek. jnT l«.ii.l992> D, trochantin fused to pleuroa (Fig. IS). Mesosternum Cartwrightand A. Ufells, NTM; D cro\ 5 9 0, 31° (Fig. %) with a pair at elongate scleritesposteriorly. 44 S 142° 29'E> Gunshot Creek, at Telegraph Legs(Figs 1&. 19) short flattened, tarai withdense Ciosving, I7.ii.l°92. Q Cartwrighl andA. Wells,QM. fringes of setae, claw on fore limb strongly curved. 11 Ofcr, 8 9 9- sameloc. andcollectors, t4-tS.iU992. Fore tarsus Mith an adpressed row ofmodified 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.. Abdomenelongate,witha lateral fringeoffinesetae, 4-5.io.1992, M. Grassland, ANIC: 31 cror, 19 9 9, wiihoutgrlis. but with a group ofanal papillae (Fig. same loc and collector. 10-U.iv.i992, NMV; I pupa, 7); anal prolegs elongate, with cluster of long setae same loc, D. Cartwright and A Wells, !8.iv1992, anteriortotheslenderstronglycurvedanalclaw (Fig. NTM; 2 larvae, same loc and collectors, NTM; 7 20). larvae U reared to aduk 9>. 1 pupa, same Joe. and Pupa (Figs9-lL 21). Length 7.5-1(13mm Labrurn collectors, 19.ii.1992. CFig. 9) short, with a small marginal papilla apJco- Neboiss(1980) illustratesthegeneral body features tnesally. Mandibles(Fig. 10), simple, slender,curved ofmaleHyatopvjchedvijancm andgivesanteriorwing Hookplates (Fig. 21)with 3-5denticles,onabdominal , lengthas6mm, butinoursamplesmaleanterior wing segmentsHl-Vin. posterior placeson segment Vonly, length rangesfrom4.3-6.2mm. Femalesconformwith developedintohand-likeprocesses, fbsteriorabdomen malesingeneral features, includingreduced 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 fragmentedsilkentubes. Theexactarchitectureofthe tentorium, is figured (Fig. l>: its anus arc well larval dwelling isunknown, butthebranchedportions developed. v>llh inner lateral flanges posteriorly, and oftubes appear to have all the features ofthe tubes lhe posterior bridge complete ofPhyhcentropus piacidus (see Wallace el at 1976) ftmale(Figs 1-3). Actetroi wuiglength6.0-8JQmm. 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 gonopodotsegmenl 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&teriorlysegmentXisdividedm formapairofdistal a blunt-ended vertical branch with a small opening lobes,each withapicalcercusandonepapilUip^TlinHy projected into the flow, a series of connected cross fusedon inner margin, thesecondprojectingveoually tunnels, and a dilated net chamber. from the outer margin at about the level ofdivision Habitat. Larval tubes were collected fronta small of segment X. streamwitharatherunstable base ofcoarse 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, limbsand anal clawsbrown-yelloworpale wherethelitter, trappedagainstlogsembedded inthe 100 A. WELLS & D. CAKTWRIGHT Figs 7-8. Hyalopsyche disjuncta Neboiss, larva: 7, whole animal, lateral view; 8, mesosternum. FSicgasle9-b1aLrsHy=al1opmsmy,cheundliessjsunoctthaerNweibsoeisisn,dipcautpead:.9y labrum; 10, mandibles; H, posterior abdomen, 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 £Z\1> _ 20 N\ ti & fofc 21 V ffrw *ti 1} 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 IMMATURES OFTHE AUSTRALIAN CADD1SFI.V m sand, was tegitining in decompose. Collections nf in the wing*; as well asothermoremajor ones, such adults were obtained atlights beside small and larger as shape of head, position of antennae, reduction of streams; the smaller streams generally were almost maxillarypalpiand formofgenitalia Thesedifferences completely shaded by thecanopies ofriparian <rees. probablyrepresentairtapomorphies Forrheparticular the larger breams were open. genera. ThewingsofHyaloprychecanbederivedfrom those o( Phyfottntropui. which closely resemble polycentropodid wings, simply by loss of several Discussion cmssveins; thoseofPhylecenmypus probably represent Coanparwws of features of larval Hyatcpsyche the pleMomorphous form. Similarities such as form disjunaa with those of Phylocentmpus pinddtm, R nf the tentorium (Figs I, 4) are probably also taddusandP. cumtinus(fromRoss &Gibbs KT73 Fig. plesiomorphous, 2; Wiggins 1977 Fig. 15.4D, and this work. Figs Superficially, the female genitalia ofHyaJopsycht ty, 15) reveal extraordinarily close similarities between disjancta(Fig. 3)appeartobestrikinglydifferentfrom species in the two genera, In derived or specialised those ofPhyloceniropusplacidw; fFig, 6} Yet upon feature*, In general form the heads are simitar close examination, similarities are apparent, (compareforexampleFigs 12. J3andPhyhcauropus particularly inthecerciandpapillae. Derivationofthe sp., Pigs 14. 15): mandibles nfall species ate stout, tfyatopsycheformfromthatnfPhytnceiurvpusrequites with rugose sculpturing dorsally, albeit on only the only simple elongation of the arrangement seen in right mandibleinH. dUjuncta; ventral sclentesdiffer Hiylocentropus, probably adapting Hyahpsyche slightly, with thesubmenhunofH. disjuAckerounded females for oviposition into sand or crevices, rather than subquadrate; the tabus imtrnus of the Investigation of the female of Dipseudopsis indfca maxillary palp orH. disjtwcui is broad and apioally McLachlanduringthisstudy, indicatedthatcmthebasis truncate rather than tapered and rounded; labia are offemalegenitaliaand wingform, Pirytocensropttsand modified to form virtually identical slender, elongate Hyalopsycht are closer than Dipscadapsis- spinnerets Thoracic features, too, show close Hyoicpsyche. resemblanceandthehighlyspecialised legsdifferonly Association or females and larvae of a species in in the exact shape ofthe specialised setae adpresscd Hyaiopsydxe has helped to stabilise the higher level totheniesalmarginoftheforetarsi. H. disftwaalacks taxonomy of this genus, placing rt in the femUy gillsbutconformswithPkyfocesttropusspecies in fbrtn Dipseudopstdae.Onthebasisofstudies reportedheTe 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 resemblethose illustratedbyRoss(1944, Ross &. Gibbs (1973) suggested on the basu crt Fig, 204)forRplacidta. Little information isavailable similarities between Phylo-crumpm and on pupae of Rkyloccntropux. Polycentropodinae that Phyloccmropus iscloseridthe Some of the above features occur in other caddis ancestral stock than other Dipseudopsinae larvae. R>r example, specialised elongate spinnerets (=Dipseudopsidae| and, given itsoccurrence in the are seen in larvae of some P&yebomyiidae and some Oligoccne Baltic ambers, they suggested t Eurasian Bcuouudae, although in these familiesthespinnerets origintor tnesubfamily (=famUy). Wehavenot found areproportionallyshorterthaninRkyloct'nJropasand evidence with which todispute this contention — the Hvalopsyche. Simpleana]claws*Jsooccurelsewhere, singlenorth-easternAustralianspeciesin Hyalopsyefie butareprobably plesiomorphous. Thehighlymodified, isundoubtedlyofOrientalorigin. Carefulphylogeoetic flattenedlimbs,theformoithemandiblesandthelarval analyses in the future will elucidate intrafamilial nicheare (bundonly in//yafryifycAe, PfjylocertJrofMis. relationships, but *l lastthefamily 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 forprovidingtheopportunity and financial Adult characters do not concur with those of the support for field work at °Heathlands** and the larvae. Noclear synaponiorphies can be recognised, surroundingJardincRiver3rcaofCape YotK,and for which islitex<eryreasonthatplacement ofthesegenera die generous help and assistance given to us by has been confused forso long. Amongadult features membersoftheSocietyduringourstayat*Heathlandsr' several smalldifferences ingeneral body featurescan D, Carrwrigbfs work wasundertakenwithbacking beobservedbetween Hyakptycheand PHytocentropus, from his family, and A. Wells with backingfrom the suchasthearrangementofsetaeonthemeta-*cutellum Northern Territory Museum of Arts and Sciences, (sec Figs2. 5) and die absenceof several crossvetns Darwin. We are grateful to both parties. 104 A. WELLS & D. CARTWRIGHT DrJ,S. WfeaveroftheUniversityofNewHampshire, comparisons. Adults ofPhylocentropusplacidus and USA, kindly provided material of Phylocentropus larvae ofPhylocentropus sp. were made available by lucidusandDipseudopsisindicafromhiscollectionfor DrJ. Morse, onloanfromClemsonUniversity, USA. References Banks, N. (1907) Descriptions ofnew Trichoptera. Proa theDipseudopsinae (Trichoptera, Polycentropodidae)./. Ent. Sot: mshington 8, 117-133. Georgia Em. Soc. 8, 312-16. Betten, C. (1909)NotesontheTrichopterainthecollection Schmid, F. (1980) "LesInsectesetArachnides du Canada, ofthe Indian Museum. Rec. Indian Mus. 3, 231-242. partie T. (Agriculture Canada). 296 pp. Gibbs, D.G. (1968) The larva, dwelling-lube and feedingof Ulmer, G. (1904) Uber Westafrikanische Trichopteren. a species of Protodipseudop.sis (Trichoptera: ZooL Anz. 28, 353-359. Dipseudopsidae). Proc. Ent. Soc, London 43, 73-79. (1915)TrichopterendesOstens,besondersvonCeylon Lestage,J.A. (1925) Notestrichopterologiques. VIII. Etude und Neu Guinea. Dtsck. Entomol. Z., 1915, 41-75. dugroupepolycentropidienetcataloguesystematiquedes Walker, F. (1852) Catalogue of the specimens of genres et especes decrits depuis 1907 dans Genera neuropterous insects in the collections of the British Nebionsiescst,oruAm.. Bu(l1l9.80A)nn.FiSrosct. rEenctoomrodl.ofBeltgh.e} 6s5u:b9f3a-m1i2l0y. WaMlulsaeceu,m.J.PBt.., 1W,oo19d2alppl.,(WL.oRn.do,n)&. Staats, A.A. (1976) Hyalopsychinae from Australia (Trichoptera: The larval dwelling-tube, capture net and food of Fblycentropodidae). Arch,furHydrobiotogie90, 357-361. Phylocemropusplacidus(Trichoptera: ft>Iycentropodidae). RosPhsy,locHe.nHtr.opu(s1(9T6r5i)choTptheera:ePvsoylcuhtoimoyniairdyae)h.iJs.toKraynsaosf WeAanvne.r,EnJt.oSm.,ol.HI.S,oc.&AMmaelriiccaky6,9,H.149(-I5n4.ms) The genus Em. Soc. 38, 398-400. Dipseudopsis from Asia (Trichoptera: Dipseudopsidae). (1944)TheCaddisFlies,orTrichoptera, ofIllinois. Wiggins, G.B. (1977) "Larvae of the North American BulL Illinois Nat. Hist. Survey23, 1-326. Caddisfly Genera (Trichoptera)". (University ofToronto &Gibbs, D.G. (1973)Thesubfamilyrelationshipsof Press: Toronto and Buffalo).