The genus Dipseudopsis Walker from Asia (Trichoptera: Dipseudopsidae)

48 Pages·1994·21.7 MB·English

JOHN S. WEAVERIII' & HANSMALICKY2
UniversityofNewHampshire, Durham, U.S.A.'BiologischeStationLunzderÖsterreichischen
AkademiederWissenschaften, LunzamSee,Austria.
THE GENUS DIPSEUDOPSISWALKER FROM ASIA
(TRICHOPTERA: DIPSEUDOPSIDAE)
Weaver,J.S.,III&H.Malicky, 1994.ThegenusDipseudopsisWa.\kerfromAsia(Trichoptera:
Dipseudopsidae).-TijdschriftvoorEntomologie 137: 95-142, figs. 1-55. [ISSN 0040-7496].
Published 15July 1994.
Thegenus DipseudopsisWalker fromAsia is revised, andaphylogenyofHydropsychoidea is
provided showing the relationships of its family taxa and the genera of Dipseudopsidae.
Presently 36 Asian species of Dipseudopsis are recognized, having a distribution including
Pakistan,India,SriLanka,Nepal,Bangladesh, Burma,Thailand,Cambodia,Vietnam,China,
Japan,Philippines,MalaysiaandIndonesia,includingSumatra,Java,Borneo,andSulawesi.Six
newspeciesaredescribed,Dipseudopsisadiaturixsp.n.,D.fintisp.n.,D. lucasisp.n.,D. mal-
aiseisp.n.,D. martynovisp.n.,andD.schmidisp.n.,andonenewsubspecies, D. robustioran-
damanensisssp. n. The identities ofseveral problematic species are resolved, with 16 species
names recognized as new juniorsynonyms (cf. species checklist p.102). D. nervosellaUlmer
stat.n. iselevatedfromformersubspeciesstatus.Newdescriptionsareprovidedforallspecies,
includingfiguresofthemalehindmesoapicaltibialspur(themodifiedspur),themalegenital-
ia, and other notable characteristics ofthe head, thoraxandwings, except two species recog-
nizedas nominadubia, D. onychophoraNavâs, and D. orientalis(Navis). Newlectotypes are
designatedforD. contortaBanks, D. morosaBanksandD. triclavataMartynov.Apictorialat-
las is provided for rapid species identification. The genus EodipseudopsisMarlier is removed
fromtheDipseudopsidae.
Correspondence:J.S.WeaverIII,DepartmentofEntomology,UniversityofNewHampshire,
Durham,NH,03824,U.S.A.
Keywords. -Asia, Dipseudopsis, Dipseudopsidae, Hydropsychoidea, phylogeny, systematica,
Trichoptera.
The genus DipseudopsisWalkerexhibits a number ified spur became the primary characteristic used to
ofcharacteristicsthatarequiteremarkableamongthe recognizespecies; e.g. this is the main distinguishing
Trichoptera. Theadults aretheonly members ofthe character in every couplet ofUlmer's (1951) keyto
suborder Annulipalpia Martynov having siphoning elevenspeciesofDipseudopsisoftheSundaIslands.
mouthparts, with the lacinia forming a proboscis, a
parallel modification of the galea in the higher The precise orientation ofthe modifiedspurmust
Lepidoptera. Sexual dimorphism is conspicuous in be made apparent for species identification, because
the genus, with the males having highly contrasting slight rotations ofthe tibia will show different per-
colourpatternsin theirwings andbodies, andthefe- spectivesofthespur. Someofthepreviousspeciesde-
males less striking in comparison. Also, males have scriptions based on the modified spur were ambig-
the mesoapicalspurofthehind tibiamodified, often uous, becausethefigureofthespurdidnotshowthe
twisted and/or branched. The function ofthe modi- remainderoftheleg,andhencetheactualorientation
fiedspurisunknown, butithasbeen usedalmostex- ofthespurwas uncertain. Thisproblemaccountsfor
clusivelyforspeciesdeterminations,totheextentthat several of the synonyms that we discovered in the
customary descriptions of the male genitalia often courseofthisstudy.Thefiguresofthespurhereinare
have been omitted from species descriptions. The usually ofthe left leg and include the adjacent apic-
conventionofusingthemodifiedspurasthebasisfor olateral spur and a portion ofthe tibiaand tarsus to
species descriptions was established by Ulmer providepropercomparisonofotherspecimensinthe
(1907b: 180): 'Bei Dipseudopsis sind die Genitalien sameperspective.
allerSpeciesganzgleich' (i.e: In Dipseudopsisthegen- Itseemssomewhatironicthatthetaxonomyofthis
italiaofallspeciesaretotallyalike). Hence, themod- eenus, which exhibits so manvfascinatine; character-
95
.
1 Sesiomorphic inrespectcotheoutgroupPhilopotamidae. 1 =Synapomorphiccharacter.
>^clliproem 0;rocoeälzleidabcsyelnitnder1=(cf0.;SIcXhmjiawdli1k9e80w)i.th articulation between tergum &sternum = 1 (cf.
ui Xwithout lateral papillae - 0; Xwith lateral papillae= 1 (cf. Schmid 1980).
ni IX normal 0; IX longcompletelysderotizedcylinder= 1 (cf.Schmid 1980).
rtneret short 0;spinner« longerthanothermouthparts= 1 (cf.Wiggins 1977).
mVili normal,undivided =0;sternumVIIIdividedintoapairoflaterallobes= 1 (cf.Schmid 1980).
: appendages2-segmcnted = 0; 1-segmented = 1 (cf.Schmid 1980).
antennaedistant =0:basesclosetogether= 1 (cf.Crichton 1957).
.inearlycvlindric.il.pretarsalclawslong=0;tarsiflattened,clawsshort= 1 (Ulmer1957,Gibbs 1968,Wiggins
at differ* from the following = 0; larva with branched tube-dwelling and filter-feeding behaviour as in
Wallace« al. 1976,Wiggins 1977).
11. Adultlabialpalpspresent=0;absent= 1 (Ulmer 1951).
:.:lt pronotum normal = 0;pronouimenlargedandwithdeepmedianfissure= 1.
dullwuh headandthoraxsetous=0; headandthoraxglabrous= 1.
:lt tcgulaknoblikc =0;tegula(aliened likeepaulets= 1.
lullwithgaleashort, vestigial = 0; modified intoproboscis= 1.
Larvalabrumexpandedandmoremembranous= 1,autapomorphyforPhilopotamidae(cf.Wiggins 1977).
jrva trochantinbroad,eitherwith bluntapicalangleortruncate= 1,autapomorphyforPsychomyiidae(cf.Wiggins
18. Ijrvawith mcsopleurallobes= 1,autapomorphyforXiphocentronidae(cf.Wiggins 1977).
larvawithtibiaeandtarsioflegsfused= 1,autapomorphyforXiphocentronidae(cf.Wiggins 1977).
20. Larvawithventralthoracicgillswhorled= 1,autapomorphyforHydropsychidae(cf.Wiggins 1977).
idcrmis with modified setae, especially on sternum VIII & IX and tergum = 1, autapomorphy for
Hydropsvchidac(cf.Wiggins 1977).
jrvaheadwithventrolateralbandsoftransverseridgesusedinstridulation= 1,autapomorphyforHydropsychidae(cf.
Wiggins 1977
.
jrvalheadandbodyflatteneddorsoventrally= 1,autapomorphyforEcnomidae(cf.Lepneva1964).
inalateral (ringedense= 1,autapomorphyforEcnomidae(cf.Lepneva 1964).
istics, should have becomeso confused at thespecies Depositories
level. IhisischieflybecauseBanks,Martynov,Navas,
and I'lmcrfrom 1905 to 1937proposeddescriptions Thedepositoriesofmaterial areabbreviatedas fol-
ol 2^ different species from Asia and the species de- lows: BMNH, Natural HistoryMuseum (formerlythe
scriptionsby Banksand Navasarepoor.Anotherfac- British MuseumofNatural History), London; BPBM,
tor might be that specimens ofDipseudopsisare not Bernice Pauahi Bishop Museum, Honolulu; CASC,
particularly abundant in museum collections, even CaliforniaAcademyofSciences,San Francisco;CNCI,
though adults areattracted to light traps. CanadianNationalCollection,Ottawa;CLDD,collec-
The primaryobjectiveol thisworkistoclarifythe tion of D. G. Denning (now deposited at CASC);
• -lieAsian speciesof Dipseudopsis. Toac- CLHM,collectionofH.Malicky;CLJW,collectionofJ.
complish this we have examined the typesofmostof S. Weaver III; MCZC, Museum of Comparative
is well as other specimens from several Zoology, Harvard University, Cambridge; MNHN,
museumsandprivatecollections.Thespeciesdescrip- Museum National d'Histoire Naturelle, Paris; MZBS,
tions have been expanded beyond the characteristics Museu de Zoologia, Barcelona; MZLU, Museum for
of the modified spur to include colouration of the Zoology, University ofLund; NHMW, Naturhistori-
1 illustrationsofthemalegenitalia, schesMuseumWien;NHRS,NaturhistoriskaRiksmu-
most ot the redescriptions ol species herein seet, Stockholm; NZSI, National Zoological Collec-
thefirst illustrationsofthemalegenitalia.We tion, Zoological Survey of India, Calcutta; RMNH,
ï Dipicudnpsis from Asia, 30 Nationaal NatuurhistorischMuseum (formerlyRijks-
ind six newspecies, and museum van Natuurlijke Historie), Leiden; SOFM,
one new subspcci recognize the names of National Museum ofNatural History, Sofia; UOPJ,
id one subspecies as new junior syno- Entomological Laboratory, University of Osaka
nyms. Two additional Asian species, D. onychophora Prefecture, Sakai; USNM, Smithsonian Institution,
and I). orientali' recognized asno- United States National MuseumofNatural History,
mina dubia, because both were poorlydescribed and Washington; ZRAS, Zoological Institute ofthe Rus-
theirtypescould notbelocated forexamination. sianAcademyofSciences,St.Petersburg;ZMAN,Zoo-
96
Weaver&Malicky: DipseudopsisfromAsia
logisch Museum, Amsterdam; ZMHB, Zoologisches DIPSEUDOPSIDAEULMER
Museum der Humboldt-Universität Berlin; ZMUC, ThefamilyDipseudopsidae is included in thesub-
ZoologiskMuseumKobenhavn;ZMUH,Zoologisches order Annulipalpia Martynov, in the infraorder
Museum der Universität Hamburg; and ZSMC, CurvipalpiaWeaver(Weaver 1984, Weaver&Morse
ZoologischeStaatssammlungMünchen. 1986). The family name, Dipseudopsinae Ulmer
(1904b), including only the nominative genus, was
METHODS first proposed as a subfamily ofthe Hydropsychidae
Curtis, butwaslaterplacedwithin thePolycentropo-
Taxonomy didae Ulmer (1906), as it has been recognized by
The morphological terminology ofwing venation several others (Tsuda 1942, Marlier 1962, Ross &
&
andgenitaliccharacters mostlyfollows thatofSchmid Gibbs 1973, Wiggins 1977, Schuster Hamilton
(1980),andSchuster&Hamilton (1984).Thespecies 1984). However, some workers in the past have re-
synonymiesarecompletelistingalloriginalspeciesde- cognizedPolycentropodinaeandtheDipseudopsinae
scriptions, butsubsequentreferenceslistedinFischer's as subfamilies ofPsychomyiidae Curtis (Ross 1956,
TrichopterorumCatalogus(1962, 1972)arenotinclu- Ross & Kingsolver 1959), and more recently, some
ded.Localitynamesincurrentuseareprovidedforspe- still recognize the Polycentropodinae as such (Flint
cimens examined, where old names taken from labels 1991).TheDipseudopsidaewerefirstgivenfullfami-
areinquotes. Forthetypedata, theterm 'holotype' is ly status by Ross (1967), and later concurred by
usedwhenthetypespecimenhasactuallybeenlabelled Gibbs(1968),Wiggins (1982),Weaver(1984),Scott
orcitedassuch, otherwiseitislistedas'type'.Anaste- (1985), Weaver & Morse (1986), and Wells &
risk(*) designatestypesofpreviouslydescribedspecies Cartwright(1993).
examinedbytheauthors. The genus Protodipseudopsis Ulmer (1909) from
theAfrotropical Region was the first addition to the
&
subfamily, a relationship later confirmed by Ross
Phylogenetics Kingsolver (1959). Two additional genera from the
Hennig86 (Farris 1988) is used for phylogenetic Afrotropical Region were placed in the Dipseudo-
analysis to determine the relationship of psinae, EodipseudopsisMarlier (1959) byoriginaldes-
Dipseudopsidae within the Hydropsychoidea s. str. ignation, and Limnoecetis Marlier (1955) originally
Weaver, and the relationships of the proposed di- placed in the Leptoceridae Leach, butlatermovedto
pseudopsid genera. Twenty four character states (ta- the Dipseudopsinae (Marlier 1961). Ross & Gibbs
ble 1) are selected, andwith polarity determined for (1973) transferred PhylocentropusBanks (1907) from
twelve taxa, using Philopotamidae Stephens as an Polycentropodinae to Dipseudopsinae, based pre-
outgroup, the data matrix (table 2) is produced. dominantly on synapomorphic larval characteristics.
These data are analyzed using the 'mhennig*' com- They recognized only three genera in the Dipseudo-
mand ofHennig86 for calculating trees. All charac- psinae, Dipseudopsis, Phylocentropus, andProtodipseu-
tershavedefaultvalues,weight= 1, activeandadditi- dopsis, and did not consider the status ofEodipseu-
ve, exceptweight= 2 forcharacter6. dopsisandLimnoecetis. However,Schmid(1980) later
Table2.DatamatrixforthefamiliesofHydropsychoideaandgeneraofDipseudopsidae,includingoutgroupPhilopotamidae.
Character
1 11111 11112 2222
12345 67890 12345 67890 1234
Taxon
Philopotamidae 00000 00000 00000 10000 0000
Psychomyiidae 11011 00000 00000 01000 0000
Xiphocentronidae 11011 00000 00000 00110 0000
Hydropsychidae 11100 10000 00000 00001 1100
Polycentropodidae 11100 11000 00000 00000 0000
Ecnomidae 11100 11000 00000 00000 0011
Dipseudopsis 11101 01111 01111 00000 0000
omo
Protodipseudopsis 11101 01111 00000 0000
Phylocentropus 11101 01111 00000 00000 0000
omo
Limnoecetis 1110? 011?? ????? ????
Hyalopsyche 11101 01111 10000 00000 0000
Hyalopsychella 11??? ?1??? 10000 ????;> ????
97
HUI VOO»
l
madethenovelassignmentofplacingPhylocentropusin that the antenna! scapes of Dipseudopsidae are not
the family Hyalopsychidae Lestage, >>n the basis ol merely close together but rather are so enlarged that
aduli characteristicsshared with thegenus Hyalopsyche theyoccupymostofthespacebetween theeyesalong
l'Ima Jvells & * artwrighi tl1)l)3> recently theanterodorsal margin ofthehead (fig. 13).
concurred with the position i>t Ross and Gibbs, and
n ilit- morphology ol the larva and temale ol Conclusions.-MonophylyoftheDipseudopsidae,
idded this genus io the Dipseudopsidae, including Dipseudopsis, Hyalopsyche, Hyalopsychella
andaccordinglyproposedthesuppressionol thefamily Ulmer (1930), Limnoecetis, Phylocentropus, and
croup name Hyalopsychidae. Our phylogeneric anal- Protodipseudopsis, is unequivocal. Two clades are rec-
es theclassificationsot Ross c\ < iibbsand Wells ognized among these genera, {Hyalopsyche + Hyalo-
trtwright, versus that ot Schmid. psychella) supported by synapomorphy 11 (cf. table
1), and {Dipseudopsis + Protodipseudopsis + Limnoe-
cetis) supported by synapomorphies 12-14. Hence,
Results
the sister group ofthe genus Dipseudopsisis unclear.
Phylogeny. - The analysis or the data matrix via TheadultofLimnoecetis, amonotypicgenusendem-
1Iennig86generated threetreeswithdifferenttopolo- ic to Lake Tanganyika, are quite remarkable, having
gies, havinglength 28, CI 0.89, Rl 0.91 (fig. 55).All uniqueadaptationstowalkonwater.Thestrikingau-
three trees depict Dipseudopsidae as a monophyletic tapomorphies of Limnoecetis present a formidable
group,comprisingsixgenera.Thefirsttreehasthefe- challenge in resolvingthe trichotomy {Dipseudopsis+
west number or ancestors, having 18 nodes, and is Protodipseudopsis+ Limnoecetis). Oneobviouspredic-
also identical to the strict consensus tree (fig. 56) of tionbasedonourphylogeny,isthattheunknownlar-
the three trees. Theconsensus tree depicts ((Psycho- val forms ofHyalopsychellaand Limnoecetisprobably
myiidae. Xiphocentronidae Ross) ((Hydropsychidae, share manysimilarities with the dipseudopsid larvae
Ecnomidae Ulmer, Polycentropodidae) Dipseudo- known. The phylogenyalso providesevidenceforre-
psidae)). thus supporting monophyly ofthe Hydro- moving Eodipseudopsisfrom the Dipseudopsidae (cf.
psychoidea. The clade comprising Hydropsychidae, finalsectionoftext).
Polycentropodidae, Ecnomidae is recognized as the
sistergroupofthe Dipseudopsidae. Afewdifferences The sister group relationship of the genus
are noted among the three original trees: 1) trees Phylocentropusremainsuncertain. Phylocentropusphe-
and 2 depict thetrichotomy(Hydropsychidae, Ecno- neticallyagreescloselywith Hyalopsyche, butthesim-
midae. Polycentropodidae),which isresolvedas (Hy- ilaritiesofthesetwogeneraarepredominantlyplesio-
dropsychidae (Ecnomidae, Polycentropodidae)) in morphic. Thus, the phylogenetic analysis abovedoes
tree I;2) trees and 1 each havea basal and adistal not provide supporting evidence for Hyalopsychidae
trichotomy among the genera of Dipseudopsidae, s. str. Schmid (1980) and in fact, infers that this tax-
both olwhich areresolved in tree2. on might be paraphyletic. Thereforewe concurwith
RossandGibbsforthemostpartontheclassification
( haracters.-Thecombinationofcharacteristicsof ofPhylocentropus, and Wells and Cartwright on the
the Iemale genitalia as exhibited in the Dipseudo- placement ofHyalopsycheand the suppression ofthe
psidae 'having synapomorphy 2, lateral papillae ol' Hyalopsychidae, ataxon thatpresentlyserves no use-
segment X present, hut lackingsynapomorphy6, ab- ful purposein phylogenetics.
dominal sternite VIII not subdivided into lateral
only a•uInneisquneotcormebpriensaetnitonanofapcohmaroarcptehriicstisctsatew,hibcuht KeystogeneraofDipseudopsidae
happens to define the family vers well, hut only in a Larvae
•lit sense. Synapomorphy 7, inferior appen- ThelarvaeofHyalopsychellaUlmerandLimnoecetis
mented is a homoplasy in Polycentro- Marlierareunknown.
and Ecnomidae. Three synapomorphies 8-
',-ort monophyly of" Dipseudopsidae, and the 1. Mandibles with apicolateral teeth; head capsule
linai hases close together, is not short, with lateral margins nearly parallel and its
unique, mbutHywdirthoipnsycthhienaIelydarnodpsyMcahcoriodncaemaistianlaseo t1o9t5a7l,leGnigbtbhsle1s9s6t8h)an itsmaximumwidth (Ulmer2
since this characteristic Mandibles without apicolateral teeth; head cap-
s absent tor the most part in Arctopsychinae and sule long, with lateral margins tapering anteriad,
Diplcctroninae. it is not part of the groundplan of andwith its length slightlygreaterthan its maxi-
Hydro; ind thus is.i homoplasy.Amoreac- mum width (Wiggins 1977,Wells &Cartwright
curatedescription ofthisconditionasahomologueis 1993) 3
98
.
Weaver&Mai.ICKY: DipseudopsisfromAsia
2. Frontoclypeal suture V-shaped with lateral lines DipseudopsodesLestage 1936: 170 (invalid, typespecies not
tapering irregularly posteriad toward the dorsal designated).
coronalsuture (Ulmer 1957)
DipseudopsisWalker The genus Dipseudopsis was proposed by Walker
- Frontoclypeal suture with the posterior portion (1852)withthedescriptionofD. capensisfromSouth
U-shaped, and the anterolateral lines nearly par- Africa and placed in the family Sericostomatidae
allel between theeyesandthen curvingmesad to Stephens, a family designation which was common
meet the coronal suture posteriad (Gibbs 1968) for caddisflies, exhibiting bizarre characteristics, that
ProtodipseudopsisUlmer did not agree with the diagnoses of other families.
3. Frontoclypealsuturesessileinrespecttoposterior LaterwhenMcLachlan (1863)addedasecondspecies
margin, posterior 1/3 nearly regular V-shaped to the genus, D. collarisfrom China, he placed Di-
and without hourglass-shaped constriction pseudopsis in the family Rhyacophilidae. Brauer
(Wells &Cartwright 1993) .. HyalopsycheUlmer (1868) placed it in the Hydropsychidae s. lat., near
- Frontoclypeal suture with short petiole joining CyrnusStephens. However, after examination ofthe
posteriormargin, posterior 1/3 moreirregularV- adultmouthparts, Ulmer(1904a, b) placedthegenus
shaped and with hourglass-shaped constriction inthePolycentropodidae,subfamilyDipseudopsinae.
(Wiggins 1977) PhylocentropusBanks Accordingly, most workers have placed the genus in
Polycentropodidae (Navâs 1913, Martynov 1935,
Schmid 1949). However, Banks (1931b) maintained
Adults
aconservativeapproach, havingneverrecognizedthe
1 Pronotum with setaceouswarts conspicuous and fullfamilystatusofPolycentropodidae, andfollowed
withoutdeep medianfissure 2 Brauer's placementofthegenus in the Hydropsychi-
- Pronotum with setaceous warts inconspicuous daes. lat.
andwithdeepmedianfissure 4
2. Labial palps present; forewing with fork I long Etymology. - Dipseudopsis:Greekdi, two;pseudos,
andsessile PhylocentropusBanks false; ops, eye; feminine. Theactual etymologyis un-
- Labial palps absent; forewing with fork I short certain.Thebasisofthisnameisuncertainalso,since
andpetiolateorabsent 3 there are no noticeable characters such aswingspots
3. Foreandhindwingswith forkI shortandpetio- present in the type species ofthe genus, D. capensis.
late HyalopsycheUlmer Perhapsitwasbasedontheappearanceoftheprono-
- ForeandhindwingswithforkI absent tum,havingthesetalareasexpandedanddividedbya
HyalopsychellaUlmer deepmedianfissure.
4. Tibial spurs 1, 2, 2; female with maxillary palps
three-segmented; one species endemic to Lake
Adult
Tanganyika LimnoecetisMarlier
- Tibial spurs 3, 4, 4; female with maxillary palps Head and thorax (figs. 12A-B, 13, 14): The head
five-segmented;AfricaandAsia 5 hasasparsenumberofshortinconspicuoussetaeand
5. Mouthpartswithmaxillamodifiedintoaprobos- isnearlyglabrous;sometimesitisglossyandverygla-
cis; hindwingwith median cell closed; malehind brous.Thesetalareasofboththeheadandthoraxare
mesoapical tibial spur modified, often twisted not raised like typical setaceous warts, but are flush
and/orbranched;AsiaandAfrica withothersurroundingsclerites. Thesetaeofthebo-
DipseudopsisWalker dy and wings are mostly short sparse and inconspi-
- Mouthparts without a proboscis; hindwingwith cuous, giving the imago a glabrous appearance. The
median cell open; male hind mesoapical tibial pronotum is raised to the same level as the head and
spurnormal;Africa ProtodipseudopsisUlmer mesonotum andisdividedbyadeep medianfurrow,
probably resulting from modifications ofthe lateral
setal areas, which have remained separate, but have
Systematicpart
become both enlarged and glabrous. The tegula are
DipseudopsisWalker, 1852 flatandsitlikeepauletsonthebasesoftheforewings.
Dipseudopsis\fy'alker, 1852:91.-Typespecies: Dipseudopsis Legs with tibial spur formula complete: 3, 4, 4. The
cvz/wwwWalker(monobasic). head,thorax,andsometimestheanteriorsegmentsof
NesopsycheMcLachlan 1866: 168.-Typespecies:Neopsyche the abdomen have patterns ofyellowish or orangish
flavisignataMcLachlan(monobasic).
EsperonaNavis 1915:397.-Typespecies:Esperonaorienta- brown contrastingwith darkbrown. Theforewingis
/«Navâs(monobasic). narrow, length ranging from 10 to 20 mm, usually
BathytinodesIwata 1927: 235.-Typespecies: Bathytinodes mostlybrownwithahyalinespotatm-cuandthear-
albusIwata(monobasic). culus, and sometimes with a pattern oftranslucent
99
hui i voor Entomologi).volumi i17,1994
bands 01 spots against a dark background, or with a u,ita (McLachlan, 1866) and D. benardi Navâs
striped pattern oi dark veins against lighter translu- (1930). SegmentIXwitharticulationbetween tergite
cabesnetntm,emIbIraanndesI;Vtaorrck Iloisngcitahriedrsshesosritlea,nIdIIpeatnidolVateaorre athnedtseterrgnuitme,anrdessetmebrlninugmaanlsoopaertnicjualwateinwilatthertalheviperwe;-
longand petiolate, and thediscal and medial cellsare anal appendages. The tergum ofsegment IXis much
The hindwing is shorter and sometimes bro- smaller than the sternum and usually overhangs seg-
ader than the (brewing being somewhat triangular ment X, and its apical margin may be triangular,
and lighter in colour, sometimes nearly translucent; rounded, or bilobed in dorsal view. The sternum of
forks II .\nó V are longand sessile, forks I, III, IVab- segmentIXisheavilysclerotizedandisthemostmas-
sent, the discal cell closed, and unlikeotherdipseud- sivefeatureofgenitalia, providingsupportforthein-
opsid genera described the median cell isclosed. The feriorappendages, thephallus,andtheventralarticu-
scent glands ofsternum V are present in the female lation of the preanal appendages. The sternum of
and absent in the male, their openings are rather in- segment IXis asimplestructureexceptforthemeso-
conspicuous and lack exterior projections. This con- superiorprocessesthat,alongwithsegmentX, forma
dition diricrs in Phylocentropusplacidus (Banks) exa- hoodthathangsoverthedorsalportionofthephallus
mined, havingthescentglandabsent in bothsexes. which is mostly membranous. Segment X is inferior
tosegmentIX, andsubequalinlengthto thephallus.
The mouthparts (fig. 12B, 13, 14) have been de- Thephallus is mostlymembranous dorsadandheav-
scribed by Ulmer (1904a), Cummings (1913) and ily sclerotized ventrad. Hence, segment X and the
Crichton (1957: fig. 53), and arequite unique, hav- phallusappeartofunctioninconcert,withsegmentX
ingthelaciniamodifiedintoaproboscis, functioning providingdorsalcoverageandsupporttothephallus.
as a siphoning tube. The two processes which form The inferior appendages are thumblike, with apices
the proboscisarenotheld togetherin preservedspec- directlydorsad, perhaps modified to holdsegmentX
imens, but curl away from each other distally. Each and the phallus together, as well as clasping the fe-
processisshapedlikehalfofalongslenderstraw,with maleduringcopulation.
its longitudinal edges fringed with numerous short
scalelikesetae,havingmanytransverseridgesalongits Female.-Femalesareusuallylighterincolourthan
shaft (thus being semiannulate), and having several themales, andasaresulttheyaremoredullandhave
mesal dentations at its base. The proboscis is at least fewer contrasting colour patterns in their wings and
as long as the first 2-4 segments of the maxillary bodies. However, the contrasting colouration ofan
palps,orusuallyaboutthesamelengthasthelabium, unidentified female specimen (fig. 2) may be excep-
includingbase plus palps. The haustellum isslender, tional. In several species, such as D. nervosa (figs. 3,
aboutaslongasthebaseofthelabium,anditsapexis 4), where the males have brown wings with large
bilobed. Ihe frontoclypeus is large and slightly bul- translucent spots, the females have striped forewings
bous, apparently well-developed to support a pump- with dark veins against a translucent background.
ing organ. Protodipseudopsis differs noticeably from Sometimes females have forewings with faded indis-
Dipscudopsh by lacking a well-developed proboscis tinct spots that are remotely similar to translucent
(fig 12(!); 11s frontoclypeus is shorter and less bul- spots in the forewings ofmales. However, the female
bousandapparentlythepumpingorgan is reduced. forewings are most often dull brown and show no
strikingcolourpatterns.
Male. - The male is dark brown, sometimes with Genitalia (figs. 15-16): Sternum VIII forms asin-
contrastingpatterns in thewingsandbody,andmore gle plate, not divided into lateral lobes as in the
striking than in the female. The mesoapical spur of Polycentropodidae. SegmentIXhasawell-sclerotized
the hind tibia is modified, often twisted like a cork- tergite, as in some species such as D. recta (fig. 15),
serewand sometimesbranched intotwoormorepro- butis moremembranous in others (fig. 16); thester-
:t is interestingtonotethatthesamespurisal- num is mostly membranous. The vaginal apparatus
so modified in l'hylocentropus auriceps (Banks), and the vulval scale are attached to the sternum of
(Schuster &: Hamilton 1984), being exceptional segment IX by sclerotized bands and membranous
amongtheotherdipseudopsids. folds. Both thevaginal apparatus andthevulvalscale
Genitalia (fig. 17): The male genitalia are blunt, aresclerotized. Thelength ofthevaginalapparatusis
generally lacking in conspicuous characteristics, and variable,beinglongin D. recta,whereitextendsante-
the processes of the genitalia are retracted and ob- riad reachingtheanteriormarginofsterniteVIII (fig.
scured by the large blunt preanal appendages. This 15), but is much shorter in other species (fig. 16).
bluntappearanceofthemaletcrminalia, especiallyin The genitalia are generally similar to those of
dry material, probably is why males sometimes have Phylocentropus (Schmid 1980, Schuster and Hamil-
been mistaken asfemales,e.g. thetypesofD.flavisig- ton 1984), except that segment IX is more heavily
100
WEAVER&MaLICKY: DipseudopsisfromAsia
sclerotized. This diagnosis is provisional, becausethe Ulmer's (1957: 303) description of the larval re-
females ofmost species remain undescribed. We ob- treat and biology ofDipseudopsisis translated as fol-
serve that the females ofsome species can be distin- lows: 'Larvae and pupae live in calm pools ofrivers,
guished from others by colour patterns ofthe body also in lakes and lake-outflows, in the mud or fine
andwings, andcharacteristicsofthegenitalia. sandy substrates; the larvae not free-living there, but
construct tubes out ofsecretions, covered with mud
etc., in the depths; these tube-dwellings are more or
Larva
less bent and twisted, elastic and flexible, but never-
The larva of Dipseudopsis was first described by theless fairly resistant; sometimes they are branched
Iwata(1927),althoughunderadifferentname.Akey into2or3 tubes, butonlyoneofthebranchesreach-
separating the larvae of Dipseudopsis and Protodi- es the [substrate] surface and there it is closed, while
pseudopsiswasprovidedbyUlmer(1957), andasimi- theotherlowerendsareopen.' Itseems obvious that
larkeywasprovided byMarlier (1962), wherebythe Ulmer's orientation of the tube-dwelling is upside
larvaeofDipseudopsisand Protodipseudopsiswere dis- down; his description should depict the two open
tinguished bythegularsclerite, mandibles, and hind ends reachingthesurface, and theone closed branch
tarsal claw. Protodipseudopsis and Dipseudopsis are completelyburied in thesubstrate. Thiswould agree
uniqueinhavingmandibleswithapicolateral teeth, a more with the tube-dwellings described in detail for
probablesynapomorphy (Ulmer 1957: f. 401; Gibbs PhylocentropusandProtodipseudopsis, andthedescrip-
1968: fig. 1;Marlier 1962: fig. 50; Marlier 1979: fig. tion by Scott (1985: 334) for Dipseudopsisin south-
5). The known larvae ofDipseudopsisknown can be ernAfrica: 'The larvae are bottom dwellers in stand-
distinguished from those ofother dipseudopsids by ing or slow-flowing water where they construct
havinghead capsule short, lateral margins nearly pa- U-shaped silk-lined tubes with a transverse net in
rallel with totallength less than its maximumwidth, mud or silt or on submerged branches, filtering out
frontoclypealsutureV-shapedandthelaterallinesta- algaeandorganicparticlesonwhich theyfeed.'
peringirregularlyandposteriadtowardthedorsalco- The larvae and pupae of Dipseudopsis inhabit a
ronalsuture, andmandibleswithapicolateral teeth. rangeoffreshwaterhabitats,includingthesandydep-
ositional zones oflakes and the pool areas ofrivers
Larval behaviour and habitat. — The larval tube- and streams. Marlier (1953, 1962) reports that im-
dwelling and feeding-behaviour have been described matures inAfricawerefound inshallowwater, while
forProtodipseudopsis(Gibbs 1968) andPhylocentropus Tsuda (1939) reported thatlarvae inJapanwere dis-
(Wallace et al. 1976, Wiggins 1977). The larvae of covered at a depth of 90 m. The collections ofD.
& m
these genera live in the bottom substrates ofstreams diehliMalicky Weaver, atelevationsof1,400 in
or lakes and construct a somewhat Y-shaped silken Sumatra, devoid of large rivers and lakes, suggests
tubecomposedofsandgrains anddetritus. Theends thatsomespeciesalso liveinmountainstreams.
ofthe two upper branches, usually one vertical and
the other inclined, are open and exposed to the
Distribution
stream current, the rest ofthe tube is located within
the bottom substrate, and the end of the bottom Dipseudopsisis known to occurgenerally through-
branchisclosed.Theopeningoftheverticalbranchis out the old world tropics ofAfrica and Asia. The
an intakeandthe inclinedbranch an outlet, through Africanfaunarecorded fromthenorthernNileRiver
whichadirectionalflowisestablishedviarapidoscil- Valley, including Egypt, Sudan, and throughout
lations ofthe larva. This forced flow enables a fine most oftheAfrotropical Region, from Ethiopia and
capturenet,withinanenlargedchamberoftheoutlet Kenya to Nigeria, Ghana and Sierra Leone, and to
branch, to collect fine organic food particles for the Zaire, Mozambique, South Africa, and Madagascar.
larva to eat. The larval tubes of Hyalopsyche are The genus is represented by42 described species, 23
branched and appear to be similar to those of fromtheAfricancontinent,and 19fromMadagascar.
&
Phylocentropus(Wells Cartwright 1993), as arethe However, in light ofthe problems discovered in this
tube-dwellings of Dipseudopsis. Therefore, since work, wesuspectthattheAfricanspecies are in need
branchedtube-dwellingsarecharacteristicforthelar- ofa thorough taxonomie revision. The Asian fauna
vae of Dipseudopsis, Hyalopsyche, Protodipseudopsis, with36recognizedspeciesisknownthroughoutmost
and Phylocentropus, this type oflarval retreat and its ofthe Oriental Region and part ofthe southeastern
associated mode of feeding-behaviour is a synapo- Palearctic Region, including Pakistan, India, Sri
morphy ofDipseudopsidae. The larva retreat ofthe Lanka, Nepal, Bangladesh, Burma, Thailand,
hydropsychidgenusMacrostemumKolenati (Wiggins Cambodia, Vietnam, China, Japan, Philippines,
1977, as Macronema Pictet) is generally similar, but Malaysia, and Indonesia, including Sumatra, Java,
differsin manydetailsandiscertainlyahomoplasy. Borneo andSulawesi.
101
.
liKui voor Entomologi!,von mi
( lui.klisi of Dipscudopsistrom Asia petersorumSchmid & Denning, 1979syn. n.
volutaWmei, 1906. p. 138
sp. n. p. 102 onyebophoraNavâs, 1935 nomen dubium p. 139
p. 104 orientalis(Navâs, 1913) nomen dubium p. 139
\.i\.iv 1930 syn. n.
Navis, 1931 syn. n. Dipseudopsisadiatwixsp. n.
uknert Schmid <S: Denning, 1979syn. n. (fig. 18)
btcolor.iuMartynov, 1935. p. 107
colleraMelachlan, 186J. p. 109 Typematerial.-Holotypea:INDONESIA:Sumatra,Huta
stelLiuMcLachlan, 1875syn. n. Padang,26.1.1990,Diehl(CLHM).
albailwjta. 192
.- Banks. 1916syn. n. Male.-Headwithdorsumandfrontoclypeusdark
discon N.i\.is. 1924 syn. n. brown, almostblack, butposteriordorsalwartsyello-
contort.! Banks, 1931b. p. 110 wish brown. Basalsegmentsofantennaebrown (type
<&A#Malicky& Weaver, 1988. p. 112 withdistalpartsofantennaeandmouthpartsmissing
.. I'lmcr. 1907a. p. 112 duetoantattack). Thorax, tergitesandsternitesdark
doehleriUlmer, 1929 (döhleri). p. 115 brown. Coxae offore and mid legs brown, coxa of
elongataBanks, 192(I p. 116 hind legs yellowish brown. Basal M of fore femora
.'...•.'.;(McLachlan, 1866) p. 116 darkbrown, femoraofmidandhindlegswithbasalA
finitisp. n. p. 116 brownish; distalAl ofallfemora, tibiaofforeandmid
immaculataLimer, 1905. p. 118 legs, andall tarsiyellow; distal Aofhindtibiabrown.
moesta Banks. 1931bsyn. n. Modifiedspur(fig. 18) longandslender,morethan'A
indicaMcLachlan, 1875. p. 120 aslongasadjacenttarsalsegment, morethan2xlong-
buddhaBanks, 1913 syn. n. erthanouterspur, distalclawabout%aslongasspur
i. sindicaMartynov, 1935 syn. n. and bent obliquely laterally, and with short lateral
infuscataMcLachlan, 1875. 120 thorn at %length from base. Forewing 16 mm, slen-
£>;,/ƒ>ƒ>/Schmid & Denning, 1979. 121 der, dark brown, but with nervation darker, having
lamellataMartynov, 1935. 121 hyaline lunula at m-cu, and indistinct translucent
M
sp. n. 123 spots between and Cu, and near apex of 1A and
4
maculataLimer, 1907a. 123 2A. Hindwing 10 mm, darkbrown, with hyalinelu-
malaiseisp. n. 124 nula at m-cu. Abdomen yellowish brown. Genitalia
»litrtynorisp. n. 124 (18A-D): Tergum IX broad and bilobate with me-
modestaBanks, 1911. 124 dian sized cavity below in dorsal view; sternum IX
pallidiMartynov, 1935 syn. n. with mesosuperior process fingerlike in lateral view.
morosaBanks, 1924. p. 125 Dorsal edgeofsegmentXwithstrongincision in lat-
nebulosaAibarda, 1881. p. 127 eral view, but apex not incised. Preanal appendages
nervosaBrauer, 1868. p. 127 longand triangular in lateral view. Inferiorappenda-
luctuosaBanks, 1913. ges curved dorsad in lateral view, with distal part
a Navis, 1931 syn. n. slightlydavateandbasolateralmesalmarginssquarish
ellalimer. 1951 stat. n. p. 128 in caudal view. Phallus calyx-like in ventral view,
ntcuwenhuisiUlmcr, 1909. p. 130 aboutYsaslongassegmentX.
iiricius, 1781). p. 130
bomt Iflmer, 1915 syn. n. Etymology. -Anoun in apposition: namedaftera
Martynov, 1935. p. 131 CeltwholivedinAustria2000yearsago.
./Martynov, 1935 syn. n. Distribution. -Indonesia: Sumatra.
robust: timer, 1929stat. n. p. 132
atkhbaiillaanSdicchamSicdh&mcidDe&nniDnegn,ni1n9g7,91s9y7n9. sny.n. uniRqeumeartkyspe..-ItThhaissasmpoedciifesieidssopnulryskimniolawrntofrD.ofmlatvh-e
junktMarlier, 19~l> syn. n. isig?iata, beinglongslender, acuminate, slightlysinu-
robustiorandamanenu ssp. n. 134 ate and bearing a short subapical lateral thorn.
sp. n. 134 However, itdiffers in havingthe male modifiedspur
spectabilisBanks, 19&31a. 134 with the lateral thorn closer to its apex, rather than
stabû ky Weaver, 1988. 135 nearthebase, the male forewings mostlydarkbrown
toni 21. 135 and without a striking pattern oftranslucent stripes
triclavataMartynov, 1935. 137 and bycharacteristicsofthemalegenitalia.
variansUlmer, 1929. 138
102
WEAVER 6cMalicky: DipseudopsisfromAsia
Pictorialatlas (p. 103-106)
Apictorialatlas isprovidedforquickspeciesdeter- riorappendage(butforsubspecies,onlyfiguresofthe
mination. The figures ofthe following male charac- modified spur). The distributional dataare listed be-
ters are provided for each species: 1) dorsal aspectof low the figures. Possible identifications should be
segmentIXandX,2)shapeandrelativesizeoftheleft confirmedwith thecompletespeciesdescriptionsand
modifiedspur, and3) caudalaspectofthe right infe- figures in thetext.
D. nebulosa Albarda D. flavisignata (McLachlan)
Burma,Thailand,
Sumatra, Malay Pen
Sulawesi
D. adiaturix Weaver
D. immaculata Ulmer and Malicky
Sumatra, Borneo, Malay Pen
D. schmidi Weaver and Malicky
India, Bangladesh
D. varions Ulmer
Thailand, Malay Pen
D. doehleri Ulmer
Assam, Burma,Thailand
India, Bangladesh
103
•KUI VOOR I NTOMOLOGU,VOLI Ml
/). collons McLachlan D. diehli Malicky and Weaver
China. Japan. Philippines
D. robustior Ulmer
D. spectabilis Banks
Burma.Thailand. Vietnam.
Borneo
Cambodia. Malay Pen.
D. knappi Schmid and Denning
D. r. andamanensis
Weaver and Malicky
Andaman Is.
D. infuscata McLachlan Thailand
D. nervosa Brauer
Sumatra. Borneo. Java. Sulawesi
Philippines
D. notata (Fab.)
India. Sri Lanka
Dipteudoptit benaniiNzvàs, 1930 1962: 5;Fischer 1972:3.
Dipseudopsis bzngana Navis, 1930: l4l, Type 9*, VIETNAM
GT.onBkeinnf'as:rudiPtheluLtLa.an-gUTlhmueo\r1n9sg13a9,00m:5e1:91d049a916t,,,adTf(icygtm.p.ne7hP5n.b)N.[amv-iissFdeitsS].c.hJe.r, DipCF'BsiTHeeosInuncNdakhAoriep:dnrs'(i:1Gsm9unP6aahh2rnn:ucg)uf.5Ll;oaa-tnnFagiFgis:NscTachhhv'euerâCorson1,n1g9g97,-612t29:1:c39h1210e.:09o;,Su8Fy'd5ines-.tcC8.hn6ae.P,nr.t1HNo9oan7lv2âo:(stm5yz.Spb.SesyJ).n,..SG*n-.,.
104