YU ISSN 0351 -0077 PRIRODOSLOVNI MUZEJ SLOVENIJE MUSEUM HISTORIAE NATURALIS SLOVENIAE SCOPOLIA 16 Botanica Ceologica & Palaeontologica Museolügica 10 Zoologica Ignac SIVEC, Bill P. STARK and Shigekazu UCHIDA: Synopsis of the World Genera of Pertinae (Plecoptera: Periidae) Pregled rodov poddružine Perlinae (Plecoptera: Periidae) sveta SCOPOL! A No 16 pp. 1 ~ 66 Ljubljana Dec. 1988 SCOPOLIA Glasilo Prirodoslovnega muzeja Slovenije. Jzc^ja Prirodoslosni muzg Slovenije, soDTiani;in
  • s and generic diagnoses are presented for adults and larvae Cüucmtfperla and SirnpUpetla are plated as synonyms of Paragnetina and Neoperh respectively. IZVLEČEK - PREGLED RODOV PODDRUŽJME PERLINAE (PLECOPTERA: PERLIDAE) SVETA - Podan je prikaz taksononiskega statusa in filogenetskih odnosov rodov poddnižine Perlinae sveta, Poddrjžino tvori 19 rodov, med katerimi sta na novo opisana Tyloperia Sivec & Stauk tn Ftircaperia Si v ec. Skupine rodov tribusa Perlini Perla /Perla, Eoperia, Dmocras), Etrotior^ma, fEtrocoretm, Kami-muria. I^eaperlopi. Tenopina) in Paro^neiitsa (Paragnaina. Tosoperh. Tytoper!a. Oyamia, Aaneiina. Marikamea} so veijelno sestrska sirupi na tribusa Neoperlini {N eoperia. Cliinopcrla. Phanoperh, Furca-perlal Rod Cfaaiisenia imamo la sestrsko skupino vseh ostalih perlin, medtem ko ost^a zaradi pomanjkljivi h pod al kov nejasen položaj rodu Aiiniperla. Preufili smo ves dostopen tipski material predstavnikov p^dTTJŽine Perltnae. Za odrasle in larval ne stadije predstavljamo diayioze rodov in do!f)äevalrie ključe. Caucasoperla is nov sinonim nadu Paragnelina, Simpliperla pa je nov sinonim rodu fteoperla. Contents U IntroiJuciiDt] .......................................................... 2. Malerial and Melhods.......................................... 3. Acknowledgements ........................ 4. Phylogeny ....................................... 5. Key tfl Perimae Males.................... 6. Key to Ptrlinac Females ..................... 7. Key to Ptrlinae Larvae........................ S, Tribus Claasseniini ............................. 8.1, C/fl«jjfnifl Wu .............................. 9. Tribus Perlini ................................. 9.\. Tyhptrla Sivec & SurSt ................. 9.2, Togoperla Klapaick ....................... 9.3, fiara^pieiina Klapakk ................. 9.4, Oyatiiia KlapaM ........................... 9.5, Agtielinü Klapäleli ......................... Manhamea Kbpaiek ..................... 9.7. Kamimuria Klapalek ..................... 9,3. EtrtKorema Klapalek ..................... 9.9. T^lropi'na Klapaiet......................... 9.10. Neop^'rlops Banks ........................... 9.11. f üeofiroy ................................ 9-12, Eoperh lllicH ................................. 9.13. Dinocras Klapalek ,..,.................... 10. Tribus Neoperlini ................................. 10.1. /-'uret^fj-ia Sivec............................. 10.2. Need ha in......................... lOJ. CkinaperSa ZwicY ........................ 10.4. Ptmneiperlae-Atils .......................... 11. Miniperta Kawai ................................................. 12. Speties not included in the lists of valid species Povzetek.......................................... Refcrcnees..................................... 3 5 5 9 10 11 12 12 14 14 17 21 23 2f> 29 32 J5 38 40 41 44 46 49 49 50 54 56 59 59 60 61 1, S'lvec. HP, Scark, S. Uchida: Synopsis of I he World Genera of Perlinae (Plecaptera: Pcrlidae) 3 I. Introdudiati At ihe time of the lu.rrs (1966) catalogue, ihinecn genera were placed in the subfaniity Pcrlinae, but few of these were well known. Much confusion existed ahoui generic limiti and similar species were described first in one genus and then in another, so that »well-known« groups like Perla. Paraf;neiinci, KciFnimiimi ami To^opcrh became almost inextricably tangled. A^ncihm Was at that time an übseure synonym of/^J>iwra,vwhile Marthamea and Phas-^anophora could not he reliably separated and EinK orema was so poorly known tliat a generic synonym, Neix'iiryptax was iisied in a separate subfamily, Acroneuriinae. Krom this abysmal state, gradual progress has been miide in pcrline systematics, particularly in the past deeade. Stark & Gaufin's( 1976) study indicated many characters of potential taxoromic and phylogenetic significance occur on eggs, internal genitalia and in nymphal provcntriculi and setatior patterns. Although this study emphasized Nearctie Perlidae, a tribal classification for the world fauna was proposed, and Claus^mia was transferred to Perlinae, Zweck tl9Büa,bjySK 1982a,b,d, 1983. 19M, 1986.198S) has made major contributions to our knowfedge of the Oriental Neoperiini and in another series of papers (ZwicK, 1982 c, e, 1984 a,b, e, 1985, 198S)he ekrified the status of Ehoforema, ienopim, NvnpeThp^., and Tyhpyge, which were formerly essentially unknown. Other studies including data on Oriental Meoperlin i during t his period are Stark & H.^L'Mann, 1978, Stark, 1983, 1987, Stark & Szczvtko, 1979, Ssvfc, i9s4, Sivf.c ZwiCK, 1987 and Zwk k & sivec. 1980. staj4k.. 1986 and Stark & Szt/V[ K.u, 19SI. include liinited data on Oriental Agm'iifta and Para^iwism in their treatment of the Nearctie members of these genera. Hari'hr (1976. 1977) included descriptions Kamimuria, }perki and S'hampeySa in his studies of Himalayan stonellies. Despite all these siudies the »Gordian knot« remained intact, particularly in refi;rencc to Kamiiijuiia. Tyhpyse. Paragnains. and Toj^opeih systematics for the Oriental regioTi-The only dear solution ED this problem lay in a massive study of museum typic material in whieh we have been involved Ibr the past decade, All available types of named Perlini species ha veno w been exainincdby us. butit has notbecnpossibletochecklbrpcrlincspecies which may have been described in other families (e. g. Perlodidae). Oti a visit to China one of us (IS) Ibund all existing Wu and Chu types are now in Peking (Zoological Institute, Academica Sinica); types named pi ior to 1962 are all destroyed and the »neotypes« named or labelled as such by Wtj (1962) have no status under the code, In this paper, we ancmpt to »unravel the knot« at least at the generic It vel, and present our current understanding of relationships an5ong these genera. Our overview oi^this fauna is necessvarily based largely on dried museum specimens collected by non-Plccopleran specialists, supplemented with fresh, alcohol preserved specimens from Japan and a few scattered n^ainland localities. Hopcfijlly this report will stimulate and facilitate a eontinufid collecting and research so badly needed on these animals, many of which are doubtlessly endangered, or extinct through habitat degradation, I. Material and Mtthtids An attempt has been made to study all available type material within subfamily Perlinae including valid and uncertain speeies recorded in the catalogues iLLltiS, 1966, Zw ilk. 1973, Specimens ofall pi iniary types and several additional ones were borrowed from the museums listed below. The most abundant material came from the authors' collections from North America, Japan and Europe. Most Onental specimens borrowed from museums were collected during light trapping for other insect groups and there is very little additional label in- formation about the eirtu m stances under which the specimens were to Heeled. Most museum speL-imtins were dry and pinred and liad to be relaxed in a moist chamber for about an hour. Male abdomens were detaehed and treated with a cold maceration technique (ZwiCK, 1983) or briefly treated with hot K OH so the penis could be removed. For a detailed study artificial eversion of the internal sac is needed. The best way to achieve this is by gently squeezing the abdominal ti p of live males to evert ihe penis from its resting position, If necessary, they were temporariSy mounted in glycerol for the study of armature. Female abdominal lips were first üoakcd in water so that the eggs could be removed, (lien [hey were usually cleared in hot KOH. Genitalia have been stored in glycerol in micro via Is or inounted in Canada balsam on transparent celluloid and as a rule attached to the specimen pin. Eggs were placed in 80% etKanol, hand cleaned with forceps and agitated in an ultrasonic cleaner for several miriLtes, Specimens were transferred to acetone, air dried, and fixed on specimen stubs with a doubfe stick tape. Specimens were gold coaled in a I funinier J F sputter coater before examination with an .A.MR 1000 SEM. Adults and larvae were studied with a Wild M 5-A stereo dissecting microscope, and line drawings have been prepared with the aid of Wild, or Leit^-Dialux drawing equipment. Throughout the tent we use the abbreviations T{Terga), S(Stema) and FCl (FoHide cell impression). Penis structures were described by using the following words for the three parts: I) basal everted »envelope«. 2) intermediate une verted i»tube pe o f he m i terga 11 obes fbu nd t h rough ou 11 ii i s gro u p (t). t he presence 0 fan e ven i y spaced row peg sensilfa set on a iow, relatively straight nymphal occipital ridge (2), and the occurence of 3 basal scleriteson thereninant orAT2,3 gi lis (3). rather than täie usual 2 found in other P er I id ae (VjChida. unpublished). Among male systellogtiathan Plecopiera, a clefl T10 occurs in all Pteronarcyidac, and in so nie members of Perlodidac. T'elloperlidae, and Chioroperlidae. Less commonly, but among all Acroneuriinae. liwperlinae and scatlened among other groups, the lOth tergum is entire and paraproets are modified as external genitalia. AltJwugh the situation is far from ccrtain, we feci (iie cleft T10 is a synapomorphy for Systellcgnalha: whether all Acroneurii-nae (and other groups) display a secondarv' loss ofthe cicli. or the plesiomorphic condilion is unclear, Specics with a clefl TiO have the remaining structure developed into a lateral pair of lohes, or hemiiergites. These structures typically contain areas of sensilla basiconica, and possibly function du ring copulation cither as hooks or probes to depress the female subgenital plate, or as )»guidcs« for the penis. The location of sensilla basiconiea on the inner basal margiiis of pcrline heniitergiies suggests, in many cases, that these provide males willi information about the everted penis position rather than details ofthe female. Other sensilla in more exposed positions are assumed to provide information on the mate. The basic hemitergite tbrm in Peltoperlidae, Period id ae, and Chloroperlidae is a broad, thin unspecialiied sheet, while I'teronarcyidae and Perlinaceath display distinctively diflbrent modifications from this basic plan. We suggest the narrow, more finger-like hcmitergiles of Perlinae represent a synapomorphy for the entire subfamily. The distribution and arrangement of peg sensilla on the occiput of perhd larvae has been discussed by StaRk & Gaumn (1976). Variable, irregular arrangements of these structures occuramong Acroneuriinae and Perlodidac species, while theorderiy association ofsensilla along a more or less straight ridge is peculiar to Perlinae. Most Neoperlini have the sensUla reduced, and exhibit a bare straight ridge; a few I'erlini also have et least some mcsal reduction ofsensilla. Several gcneric clusters have been identified within Perhni on the basis of our limited data (Table 1, Fig- IV Tliree principally European genera. Pcrfa. lHoperh and Dircfirai are apparently descended from an early Kcirnimuriu üt Firm t/renia-hks ancestor with relatively small hem i tergiies and simple abdominal terga; these genera are allied by a divided anterior margin ol"T9 {27>, In other Perlinae and Acroneuriinae, the sclerotized anterior margin of T9 15 entire, but in these genera, the sclerite is interrupted in the niesal third by niembrane. Table I, Apomorphit; characters (plesiomorphy) ]. Spetrialired hemiletijites (simple DiawgMin - Cukus type) 2. Larval oKCipiEal sensiJIa roiv and ridge regylar and Complete [inlerrupled, irregular) 3. Three baial xlerites on the remnant of AT2, 3 gilli (two) 4. SeCtradary loss of abdominal pleural brushes (presencel 5. Female Jiiterstemnl 9-[0 seta I-1 ike microtrichia (small trlansular microtrichia) 6. Hammer loss (presence) 7. Sensilla basiconica cluster on heniiierjsite ape.x (absenct) 8. Para^nelma type basal penial sclerite (no basal sclerito) 9. Upturned, swollen, basal t;alluson hemitet^ite {no basal tallus) [0. T9 sclcrotizeij apterolaterally, meinhranoui mesally (T9 wilhoui anterolateral sclerite) 11, Penis sac strongly recuri,-ed, complexly lobed (penis sac straighttr. simply lobed) 12, T5 modidecj With lobe or patch :T5 unmodified) 13, Tongue-hkc, long subgcnital plate (subgenital plate short or u[l^peciali^ed^ 14, Basal lobe on penis tube (penis tube without basal lobe) 15, Small, mesocauclal lobe on T8 (TS without mesoc:audal lobe) 16, t'ompleily lobcd anterior process of hemiterjiies (simple anterior hemitergite process) 17, Anterior chorionic fold (no anterior chorionic fold) 18, Hcmiiergites with basolateral knob (no basolateral knob) 19, Slender, mesal sclerite on T9 (19 sclerite absent or of other type) 20, Hcmitergite anterior procejs with .(lender ventral and enlarged dorsal lobe (hemiicrgite atirerior process unlobed) 21, Two ocelli (three ocelli) 22, Sclerotizcd tucsocaudal process on T9 (T^ without such a process) 23, Eloiisatc, posteriorly located tentorial oallosilieiš (oval caJJosities. tbrviard of ocelli) 24, Massive subgenital plate (urispeciali7X'd subgenital plate) 25, .Anterior T9 sclerite prolonggil mesal I y [T4 anterior sclerite normal) 26, E^ anchor rnu Iti fibrous with lateral projections (egg anchor simple) 27, T9 anterior sclerite interrupted mcsally (T9 anterior sclerite complete) 28, l,arval submenium with bajolateral sutures (SLbmetitura without basolateral sutiires) 29, T9 mesal field with chit i nous spines (T9 mesal field simple) 30, Penial sac armature long and setaTikf (penial sac armature short and triangular) 31, Eeg collar rim thick with few thick ribs (collar rim thin with multiple ribs) 32, Male abdominal SS-9 with sensilla basiconica (abdominal sterna without «nsilla hasiconica) Penial tube basal icieritc with constricted opening [basal nclerite opening wide) 34. Hemittfgite baiat piece llutteiied beyond origin of anterior process (hemiter^itc without flattened basal piece} 35. Egg choriun striate (e^ chorion non-striate) 36. Secondary loss of anterior oeetlus (three ocdli} .17, Bifurcati; anterior hemitergal process (simple anterior process) .Ig. Long basyl ccrcal segment (short basal cercal segment) 39. Large black penial sac armature (brown variable sac armature) 40. Miiditied T7 (unmodified 17) 41. Slender, cleft mtsal sclerite on TS (TS without such a sclerile) 42, Short Sc vein (normal Sc vein) 43, Basally curved Cu2 vein (straight Cij2 vein) 44, Nymphal thoracic sterna with seta I fringes (ihotucic sterna without setal fringes) Wiihin Ihis eluiter, Eoperla and Dhutcrm larvae exhibit a basal suture (38) on ihe lateral submental lobes. This character is incompletely expressed in Phunoperia and probably rep resen Ls a convergence in this group. This presumed apomorphy unites these two genera, while the Perla species all exhibit unique, chili nous plates in the mesal field of T9 (29), D'mocrtss J SivK, B.P. Stark, S. Uchida; Synapsis of the World Genera of Perlinac (Plccoptcra; Ferlidae) 1 PurognMlmi S Phonepwia Fig 1. ProposM) phylogeny ofPerlinae gsnera. [Miniptrk not inclLdcd). The numbers (1-44) refer to Ubk 1. species share a sirnilfir, unusual collar form (31), und scattered sensilla ba^iconica on male abdominal fitema (32), whik Eopt'rla is distinguished by peiiial armature which Lncludcs long seta-like spines on the sac (30). Tügopcrta, Pam^^nL'lina, MarihariKti, Agnetina, and Oyamia all have a highly modified male abdominal T5 (!2). Typically this modification takes the form of a prolonged and sometimes clcfl lobe, but lobe reduction to small knobs orsensiüa basiconiua patches has apparent)' oecured independently in three I'amgnctim and an Agnetina species. !>Aiir!hümea, ARnelina and Oyamia appear to be allied by their unusually massive and complexly I obed hemitergites (1 fij, but the Togopcda-Ptiiuffncfina grouping rests entirely on an intuitive, probably plesiomorphie, similarity. Oyamia species are distinctive in the peculiarly massive form ofthe hemitergitcs which have a low lateral knob (18), \tanha7nea and Agm'lina are allied by a common chorionic fold, collar type and anchor (17), but an alternative arrangement linking Agnfiina and Oyamia on the basis of reduced penis armature (plesiomorphic?) or bilobed T6 (mis.sing in some species of both genera) is less satisfactory, Agiwiina species arc united by the iicndermssa} sdcTite on T) (^ and by fhc irmch clongafed and .^icleroti^ed hemitergites, while Marfhainea species share bilobed hemitergites with a slender ventral lobe and an expanded dorsal lobe (20). Faragm'titia species are allied by the shared process on T8 (15), similarities in e^ collar, and vaginal armature, while Togopcrla species share a distinctive subgenital plate shape (13) and a common basal lobe on the penis tube (14). Tylo-pcila is thought to be allied to the Paragnetina complex of genera on the basis of a similar, upturned basal calljs (9) and by the similar sekroti^ation pattern of T9 (10). The T5 lobe may have been independently lost in Tyioperla but we are, instead, presuming this to be due to plesiomorphy; the genus is deli ned on the basis ofthe complexly lobed and recurved penis sac (11}- Mii}iper!a probably should he included in this complex as a diminutive Agnclina - like genus, but our knowledge of this unique form is limited. The EiFOCoremu-Tetfopim-Neoperlop^ trichotomy is based on a common bioceilate condition (21). The presence ofi ocelli occurs sporadically among several Plecoptera families and presumably is the result of independent loss of the anterior ocellus in all these cases. Among Perlinae there are 2 apparent e,\amples of independent loss ol" this ocellus viz the Neoperlinigenera, Neoperia, Chinoperia, and Phunüperla^ and the Eirocorema generic complex. The position of Kamimuricf relative to this complex is tenuous since no apumorphy could be identilled, however, Kamirmina shares the same type of basal penial tube scleriie (S) with the Etroiorema and Paragnt'Sina complexes whieh suggests its placement in this grouping rather than with the Fcrla complex in which this character is lacking. Tesmpina and Seoperhps diT^ united by the unusually long and posteriorly placed tentorial callosities (23), but apomorphies have not been identified for either of these genera at present. Etrocorctna is recognized by the massive subgenital plate (24). distinctive T9 sclcritc in males (25) and by the unusual anchor on the egg (26), Kamimuria species share a sclerotized, mesocaudal projection on male T9 (22). The tribe Perlini (Consisting af Puragnefina, E^frtfiorema and Pcria complexes) is united by the presence of sensilla bas icon iea on the hemitergite apex (7), and Neoperlini, the presumptive sister group, shares secondary hammer toss with this group (6). C/ciay^ienia, the apparent sister group of all other Perlinae, has lost the pleural brushes on abdominal segments (4) which occur in essentially all other Systcllognatha, and has an unusual expression of long seta-like spines on the intersegmental membrane ofS9-10 in females (5), Neoperlini genera are united by a distinctive, complex basal penial tube sclerite. with a constricted opening (33), and by the hemitergite form, which includes an anterior process arising from a flattened basal pieoc (34). Weopcrla, Phanefpci la and Chinopcrla ai-e united by the striate eggs (3,i) and a secondary loss ofthe anterior ocellus (36). while Furcaper/a, with bifurcate anterior hemitergite processes (37) and an elongate basal cereal segment (38) stands as the sister jp-oup of this Complex. Cliinoperia and Phanopcrlu are united on the basis ol' 1. Sivec, ß.P, Stark, S. Uchtda; Synopsis of (he WorJd Gs:ncra oC Pttlinae (PlütoplCrd: PcHidne) 9 similar large, shining black penial sac armature (39), Neoperia is ü large, L-omplex genus but the greal majority or^pecieü share a similarly mod iticd T7 (40). Chinoperla species arc united by the long, slender, dert scJeritt on T8 (41) and by the very jhon Sc vein in the fore wing (42). whiJe Phanoperla specics share sytiapomorptiies in iht ba!,ally cjrved Cu2 vein (43) and the nymphal posterior thoracic sternal fnnges (44), otherwise known only among a few pel-to perl id genera. 5. Ktj to Ptrlinae MsiJes 1, Lobes or sen si I la basiconlca patches on T 5 .................................................................................- 2 T 5 ivithaul i obes or ^nsilla basiconica......................................................................................... 6 2, Hemitereites ^vith 2 or 3 lobes i it lateral asptci, or esiending Ibnv ard to at least T 8 ................ 3 Hernitei^ites with only a basal callus, unlobed ifl liiternl aspect, and reaching only T*) ............. 5 3, T 7 w ith dense m esa I sc ns il 1 a basiconica patch: hemit erga! lobes bi 1 obed i n 1 a ten! I aspect, nd beari njj a basiil callus ............................................................................................................... Manhamea T 7 with at most a few scaii.tr«i senčil la basiconica. not fonninfi a dense mesal patch; hemitcreal lobes simple, bilobcd nr trilobed in lateral aspect, but without a basal callus ............................ 4 4, T9 completely membran aus inesally.................................................................................. Oyamia T9 CO with a sclerotizcd mcs;il band ................................................................................Agne!'ma .S. Mesal fitld of TS usually sclerotiztxi and produced posieriorly into a small lobe .............................................................................................................................. Pafo^retiria Mcsal field of TS membranous and without j posterior [übt ........................................ TofiopL'rIa 6. Hiocdlate ....................................................................................................................................... 7 Triacclliite..................................................................................................................................... II 7. Hemiterignes simple, linger-like, without bflsal caJlus or ynterior process sxtendJn^ frora basal piece Hcmitergites bifurcate, with anterior linger-like proccss eütending from mcsalty direaed hasal piece ......................................................................................................................................................... 9 5, Anterior sclerotixed marigi n ofT^J projects into mesal membranous field, partially dividing tcilguni; hemitcrgites extend lor^hard over T9 ............................................................................ ^:lrtk■[)rf^?[ö Anterior mEirgin T9 straight, not project ing caudal I y; licmitcrgltes short, scarcely reach in g posterior mai^sin 79......................................................................................................................... Tetropina y, T9 with strongly sclerotizcd, usually Y-shaped mtdiat sclerite.................................... Chiiu>pcr/a T9 without sclerotizcd, Y-shaped sclerite ................................................................................... 10 10. Cu5 of fore wing curved basa 11 y, arising from Cu] al an almost right angle; T 7 without lobe .......................................................................................................................... Phanopcrici (in part) Cu2 of forewins straight basally, aristng frcim Cul at an acute anfile: T7 typically with a lobe , .............................................................................................................................................,\'eopvrlü 11. T4-5 wit h a nterior scleroti zed band and me m bra nou s posterior field: mesa! fieldofTSwit hout spi n y projections ...................................................................................................................... Minipi-rici T4-5 unifoi mly sclcroiized or mesal field cirT9 with spiny projections ................................... \1 12. HemitergitC-s bifurcate, apes of anitriorly projecting portion without sensilla basiconica ........ 13 Kemitecgiles simple, with at most a basal calius ........................................................................ 14 13. Projecting portion of hcmitergitcs forked ....................................................................... Fimapfria Projecting portion of hemiter^tes simple ........................................................ Phiitsoiierh (in pan) !4. T^) with mcsal lldd of spiny projections ....................... ........................................................ Perla f? without spiny projections in mcsal field................................................................................. 15 15. TS wlih mesal lobe and TV with mtiial mumbmnous ficM; licmitcr^ilcs with well developed basal callus.................................................................................................................................. Tvhper/a TS without lobe, TV mesal held suleroti^ed and usually covered with sensilla basiconica....... Ifi Ifi. Sclerotizcd anterinr mat^n projecting deeply into mtsal field of T9, European and North African species ................................................................................................................................................................................................................17 Sclerotizcd anterior margin of T9 straight; Asian speoiej............................................ A'arnimiiriii 17. Project ing portion of hcmitergitcs about as long as wide ................................................... EojK'ria Projecting portion of Vitmitergites longer than wtde ......................................................... fMniKfu^ 6. Key to Per) i na« Femalesi I ßiocdbtü ......................................................................................................................................... 2 Triocelblc........................................................................................................................................ 7 2. Sc not esiendiTig beyond basil ihird of forcwing ....................................................... Ctiinnperla Sc qxicnding [0 distal hairot'forewin^........................................................................................... 3 3. Subgenital plale narrowly noKhwi and massive, covering niosi of ventral alxtotninal apcK ......... ........................................................................................................................................ Klfticorcma Sut^mtd] plate small or unproduced, wiili or i^ithout notch..................................................... 4 4. Cu2 of forewing curved ba?a!ty, arising from Cul at an almoal right an^le... Phcinopcrltt [in pari) Cu2 of fore wing straight basal ly. arising from Cu! ai an acutc angle.......................................... 5 5. Tentorial callosities adjacetiL to ocelli, linear and approxiniüting ocelli in width ....................... 6 Tentorial callositit-s forward of ocelli, oval and smaller than ocelli .................................Neoperltt 6. Occiput with a small mesal knob; e^ with a dislinel ecloNion line below raicropylar line............ ........................................................................................................................................... Tetropina Oeeiput without mesaJ l:nob; egg without edosion line.............................................. NeopcrhpH I. C\i2 of forewing ciir\ed basal I y, arising from Cul at an alfnosi righl ai^gle.,. Phanoperlü (in pad) Cu2 of fo new ing straight, arising from Cul at an acute angle........................„„,„„„................... 8. Hotly length less than 10 mra; endemic to Japan ........................................................... \Jifiipcrla ftody lengtti greater than 10 itim ................................................................................................... 9 9. In te reegmen tal membra ne of S9-10 with a den se band of long setae; egg sp i nd ie sh aped wi t h button collar ................................................................................................................................Clacssi'ma In te rsegmental me tnbrane of S9-10 w il hou 1 dense band of I on ^ setae; e^ sha pe a nd col lar variable ....................................................................................................................................................... 10 10. European, or/and circum-Mediterranean species ....................................................................... 11 Asian and Nearciic species.......................................................................................................... 15 II. Subsenityl plate unproduced .................................................................................................... 12 Subsfitilal plate dislinetiy produced............................................................................................ 1,3 12. Egg with wide collar: vagina without anterior accessory glands ...........................,.,.......... t'o/vW« Egg with button-Iike attachment of anchor, vagina with aitterior 4ci.fssor^' glands........................ ............................................................................................................................Mtinhamen (in pan) 13. Subgenital plale distinctly notched; chorion granular to ornate ............................................ Pcrin Subgenital plate without notch; chorion smooth ........................................................................ 14 14. An tcrior vagi na I accessory glan ds p resen t; egg wit h bu tt on-1 i ke attac hnn en t of a n c h or, rare or ext i net poiamon species, recent records of A^nentja only from Poland. Anatolia and the Caucasus........ ......................................................................................................... Manhamm (in part), Aj^nctina A n le rio r vagi nil I accessory' gland s absen 1; egg w i th w ide. sessi 1 e CO 11 ar: com mo ti spec tes in smal I and large streams throughout mucli of Europe........................................................................ Dituxras 15. Anterior vaginal accessory glands present................................................................................... 16 Anterior vaginal accessory glands absent ................................................................................... IS 16. Subgenttal plate long, tongue shaped, usually without notch; egg chorion smooth; collar senile or absent ................................................................................................................................ TDj^operia S utonit a I plate lengtli variable, usunlly short and triangutar, often notched, egg chorion variable, collar stalked or but Ion-I ike......................................................................................................„„. 17 17. Subgenttal plate entire or truncate; vaginal scliirites present; egg collar butlon-like .......Agnijimi Subgenital plate notched, vaginal sclcntes usually absent; egg collar stalked ............ Pums^mtina 18. Spermatheca an irregular, tar^, inflated sac............................................................................... 19 - Spermatheca a slender ^tJsage-like sac....................................................................................... 20 19. Siibgenital plate reaches posterior third of S8 .................................................................. Tytoperia Subgenital plate not reaching posterior third of S8 ..................................................... Kamimittia 20. Subgenital plate short, broadly rounded and shallowly notched; basal cercal segment about 2 times long as wide; known from South China .........................................................................FtirL-apci ia Subsenita[ plate entire or truncate; basal cercal scgnicnt wider than long: known from Japan. Koixa, northeastern China and Soviet Far East ............................................................................ Ominiu I. Sivec, BP. Slaric, S. Uchidar Synopsis of the World Genera of Perl in ae (Plecoptera: Perlidae] 11 7- Key to Pcrlinae l.arvac [Furcaperla. Miniperla. Neoperhps and Chiiiapfrla unknown) !. European and circum-Mediicrraiißin species ............................................................................... 2 Asian and Nearclic spictes ............................................................................................................ ft 2. PoiEcrior seta I Iringp complete on S7 ........................................................................................,.,. } I'oiterior setnl Iritige incompfete on S7 ......................................................................................... S 3. Abdominal leina »'iih black cloihing h.iirs; lateral submental lobes without ijasal suture............. ............................................................................................................................... Agnefitia {in pan) Abdominal terga without black clothing haira; lateral submental lobes with basal :)uiures ....... 4 4. Head between etdjsiai suture and o(;cipital seta J row mostly pale; abdominal terga witti while clotEiifij; hairs; abdomen wiili 2 more or less complete pale lon^iuclinal hands............... Kopi'Ha Head between eedysial suture and occipital seta I row mostly dark; abdomii^al re]-ga with short, thick intercalary setae; abdomen without pale longitudinal bands ...........................................Dtiiixras 5. Area between posterior otelli with at least a small arey of pale pigmentation; cerci typically with dorsal swimming hairs along basal segments; posterior tergal fringes with only short thick setae ...................................................................................................................................................Peria Area between posterior ocelli entirely dark; cerci without swimming tiairs; posterior tergal fringes typically with a few long setae mised witli shoit thick ones........................................Uartkamca 6. Bioceilaic ........................................................................................................................................ 7 Triocdlate............................................................................................................................... 10 7. AntL-Folaterdl sub mental lobe:; olfet by hasal suture; one pair PSC3 gills; püstcrior margin of mesthitemum with SL'lal fring;! ...................................................................... I'hantiperia (\n part) Anttroliiteral submeni:i1 lobes withom hiavil suture; two pLliI^ PSC5 gills: pciaierioir margin of rrteso^ternum without setal fringe .........................-..................................................................... 8 a. Otcipitiil ridge with close-set. ctimplete row of short, ihick setae; body den sie I y covered with black ctolhing liairs ............................................................................................................... Telmpina Occipital ridge with at mo!jt a few shon, thick setae laterally; clothing hairs sparse iind typically brown .......................................................................................................................................... 9 •). Lateral margins of pronotum completely frin^scd with thick sstac; anal gills absent ..................... ................................................................................................................................... EirtKWi'tnti Lateral marjjins of pronotüm wilh fringe iticotriplcle; ansil gills typically present .......... Stoptrlci 10- Anterolateral suhmi:ntal lob» olVset by basal suture; one pair PSC^ gill^; ptwtcrior margin of mesosternum wilh setal fringe ..................................................................... Phuiiojierla tin part) Anterolateral submental lobw without basal suture: two pairs gills; p^Kterior niar^n of tnesostemum without setal fringe ................................................................................................ II ! 1- Anal gills present......................................................................................................................... 12 Anal gills absent ........................................................................................................................... 15 11- Posterior setal fringe ol ST incomplete ........................................................................ fawK'it'fcia Posterior setal fringe of S7 complete ............................................................................................ 1.^ 13, Posterior fringe ofT2-4 with at least a tew conspicuously long setae intermixed with short, thick ones; less than 10 thick intercalary^ setae on T9 .......................................................................... 14 Posterior fringe of consists of relatively uniforin setae; typically motiethan lOthicls intercalary setae on T9 ......................................................................................................................Ckmssi-nhi 14, Posterior fringe of metan ot urn and most abdominal lerga with multiple long setae intermixed with short thick ones ..................................................................................................................Aancliiiu Posieri or fri nge of met a not ttm and most abdomi nal te cga wit h only 1 or 2 I on ^ seta eintermixedwith short, thick ones ................................................................................................................... Oyamiu 15, Thorax and abdomen with a median row of long silky setae; proventriculus amied with two well developed groups of scle rot i zed bands ........................................................................................ 16 Thorax and abdomen without a median row of long silky setae; proventriculus with anterior group of scli:noticed h^nds reiluced and posterior group well developed .................................. Tofiopcrla 16, PSfl. 3 eill^ single ............................................................................................................ TyUiperIa PSC2. 3 gills double ........................;,................................................... ['aragneSsna, Kamimuria H. Inbus CLAASSEN[INI, new tribe 8,1. Ciaassenia WL' Adelmgia KlapAlek (1914). Type species Adelunsia laadata ECiapAlek, orig, desig. PreoccupieJ (Me-uchar, 1902). Chassenia Wu (1934). Replacemeni for Adeimgia Klafälek. Aduli habitus. - TrioceKate, large and generally dark brown. Wings greyish, veins dark brown. Females and males macropterous or brachypterous. Mesa! occipital knob abstnl. Claassenia Fig. 2. Ciaassema sabuSosa (Ba^k^) [USA]; Abdominal tip of female (a), vagina (b), head and prot haras (1); Ciaassenia radiata (KlapALek) (China); Abdominal tip of male, ventral (d), dorsal (c), and everted penis (e). I. Sivec, B.P. Stark, S. Uchida: Synopsis of the WmU Genera of PerJinae (Pleeoptera; Periidafil ! 3 Claassenia Fig. 3. Claassenia jp, [China]: Nymph habitus. Male genitalia, -T5-9 unmodified; T9 with large mesal palth ofbensilla basiconica; T8 someiimM with scauered sensillabasiconiai. Httnnergites simple, finger-like, typically reaching posterior margin of T9, at most anterior third of T9; a lew sensilia basiconica scattcn;d along inner, dorsaJ surface of hemitergite. S'J with circular hammer; abdommal sterna without brushes, Penisenvelopertiemb ranous, t ube and sac me m bran ous, un armed or wc ak ly arm ed with minute triangular spines and slender seta-like spines. Female gemtalia. - Subgenital plate weakly produced or unproduced. Intersegmental membrane of S9-10 wilh dense band of long setae. Vagina membranous, accessory glands present or absent. Spermathecal 5talk short; spermatheca targe, bulbous, with an irregular shape. Egg. - Spindle shaped with or without button-like collar. Chorion smooth or with weak FCIs on posterior pole, Micropyle? subequalorial; orifices sessile; eanals slanted, Larva, - Distinctly patterned in brown and yellow. Occipital seta! row with a few setae missing from ridge behind ocelli. Lateral margins of pronotum fringed with short thick setae; fringe sometimes incomplete, Abdominal tergal fringes uniform. Black clothing hairs present. S7 with posterior fringe complete. Cerca! swimming hairs absent. Ana J gills present. Distribution.- Nearctic, Soviet Far East. Korea, mainland Southeast Asia. Vafid species,-bishuffi (Wu) hfachyptara Bjjinl'k caudüta (klapalek) ßsh-a'^v * ifi^ai (Ki,a.pA lek) hnaisi^ia Wi; rtfüj^ffja Wu * inatjcimrians (bamks) * todima (klapALEK) * ia/j|ifojifl (BAMiS) * semihrachypiera Wu &. ClaaSSEN UiKta (NavAs) Discussion. - Several of t he Oriental-eastern Palearctic species cannot be distinguished presently and it is probable that the list above includes a number of synonyms. 9, Tribus PERUNI 9.1. Tyhperta Sivtx & St.ark, n. sen. Type species. - Tyhipygif iisii-nmta Wu $l Q,aassen= Tylopeda menuaia. here designatfK), Adult habitu.s. - Triocellate, medium to large, brown species. Wings suffused with brown, veins dark brown, Macrtiptctous, Males with dense brush of setae on anterior me-tasEernum, Male genitalia, ^T5-7 unmodified. T8 typically with a small mesal lobe covered with sensilla basiconica; lobe absent in some species, T9 sclerotizcd laterally and along anterior margin, large membranous area mesally; membrane of T9 with a small mesal patch of sen-silla basiconica. 1 lemitergitc simple, fmgcr-like, usually acute apically and usually with a well developed basal callus covered with sensilla basiconica. S4~7 with hair brushes; brushes longest on S6. Penis envelope sparsely covered with min uie, triangular spines; tube membranous 1. SivK. O P. Stark, S, Uch^da: Synopas of (he World Genera of Perlifiae (Plcwptera; PerHdae) 15 \ A V Tyloperfa Fit. 4, Tylppi'rla Jbrmosana (OkamotoI [Taiwan]: Ab<3orainal tip of female (a), abtlcuninal tip of maie tb), hemitergile (c), evened penis (d>, bead arid pTothora* (f). and tgg (e). unarmed, with apical pair of lobes; basal scleriteu well developed; sac apically recurvcd, with pair of tiasal lobes; sac armature of variable size spines covering lobes, and usually apex., Female genitalia. - Subgenital piaie large, sub triangular covering most of S9, and notched apically. Vagina membranous, acccssory glands absent. Spermathecal stalk säender, sub-equal to vaginal lenght; spemiatheca inflated, of somewhat irregular elongate shape. Egg. - Elongate oval with short wide toUar. Rim smooth, imflanged, Chorion smooth, M ic ropy les in posterior third; orifiLCS sessile. Larva. - Brown without distinctive pattern. Oeeipiial setal row complete. Pronotum completely fringed with short, thitk setae. Abdominal tergal fringes with few long setae. S7 posterior fringe ineomplele. A.nai gills absent. Cereal swimming hairs present; siiky hairs on dorsum of head, thorax and abdomen. Distribution. - Taiwan and mainland Southeast Asia, Tyloperia Fig. 5, Tyloperia Jbrmosam (Oka.wütoJ [Taiwan]; Nytnpti habitus. I. Sveč, BP. Stark, S. Uthida: Synopsis of the World Genera of Perlina^ (Pletoplera: Ptertidac) 17 ValiJ ^ipecics.- * (iiienmici tWu CuASStN), n, comb. * fitrmnänu fOfiAMCJTo), n. com. = minor (KlapAlek), siftnata n. syn. * ittnlcri (NavAs), ii. comb, Discussion. - This genus includes the distiticiive re in nan ts of Tyiopyge which was based on T. planiiiono, a species orFamgnetina (ZwjcK, 19&8). T. attc-rtuaia types are lost, but two males from China, cnllecicd by D. C Graham in July and August, are in the United States National Museum and these npecimens agret closely with Wu & Cl,AASSErv'i (1934) description and fi&ures, particularly in hemitergal ühape and tcr^f modifications. 9.2. To^oper/a K:r,AP\t,f,K Frrh (Tot?oper/a) KlapALek tlMT). Type species Pt^r/a Imftiiia Pectet, desig. KlapAlek (1923). Adult habitus. -Triocellate, large, brown to dark brown species. Wings suffused, veins brown. Macropterous. Mesa] occipital knob absent. Male genitalia. - T5 produced into a slightly excavated or hi I obed process, T 6-9 sc lernt i zed lateral ly and along anterior margin; membranous posteromesally; membrane of typically setose. Hemitcrgitcs with short, broad to slender anterior process, and well developed basal callus; ape>i of anterior process and basal calius with scnsillabasiconica. S4-8 and ntctasternum with hair brush. Penis envelope membranous; tube membranous with basal, and usually a pair of small lateral iobes; distally tube is armed with small triangular spines; sac subequal to tube and armed basally with small spines, apically with patch of long slender seta-like spines. Female genitalia, -Subgenital plate long, produced over most orS9, subtriangular to tongue-shaped and sometimes excavated or slightly notchcd apically. Vagina long, membranous with accessory glands on anterior margin. Sperm at h eca I stalk shorter than vagina; sper-matheca sausage-1 ike, often wrinkled, pointed apically and bearing a few accessory glands. Egg. - Elongate oval to elfiptica], with or without collar. Chorion smooth. Collar, when present, sessik, wide with smooth, un Hanged rim; row of pits around collar base, Micropyles in posterior third; orifices sesstic; canals slanted. Anchor, w hen present, m ushroom-like with small flat cap and short pedicel. Larva. -Mostly brown with subtle pattern of yellow on head and thoras Occipital setal row complete. Pronotum with complete fringe of short thick setae. Abdominal tergal fringes of mined long and short setae; often with mesal pair of prominent tergal intcrcalary setae. S7 posterior fringe incomplete. Black clothing hairs present. Cereal swimming hair^ present. .^llal gills absent. Distribution. - Mainland Southeast .-\sia and Japan. VaJid Bpccii,- * Jonimafi (NavA!!), n, comb, = (am7iin/?aia (Endlrlhin), grafiami Banks, n. syn. * Umbata (PirrET) = kawamuraf (Okamoto), mdt-mmura? (Okamoto), n. syn. * perpkia Klapalek = l:ifo\'eperla ptrpkia (KlapAlek) [CTiina]: Abdominal tip ortemale (a), vaj^inü [b,i;). abdnfiiinal lip of male {d>, hemitereiie (e,!). l>asa! callus of hemitergites(g)e^'ert«i penis (h>, and head and prothora* (i). J. Sivec. D.P. Stark, S, Uchida: Synopiiis of the World Genera d Perlinae (Plecoptera; PcrlEdae) 19 Togoperla Fig. 7. Togoperh iimhiira (Pjctft) pa pan]; Nyniph habius. The genitalia of the T. tonkinmsi^ (NavAs) t>'pe is missing, since no figure of the genitalia exists in the literature, this specieü is unrecognizable. Perla icntiina Nee d ham has been included in Togoperia since KlapAlek-b (1907) study but it is a m ember of AflVM/wi (Periodi dae). Fig. 8. Puragneiina lacrimosa (Ki apAlek) [China]: Abdomina! tip of female (a), vagina (b), abdominal tip of male (c), hcmitergitc (d), ewrted petiis (c3. and veniral view of everted sac (f) i. Sivec, B.P. Stark, S. L'chida: Synopsis of the World Genera of Perliiwe (Ptecoptera; Periidae) 21 J. ParagRetina kLAPÄLKk Peria {Famaneiina) KlapAlek (1*^07). Type species Perta mil'spennm McLachlan, monotypy. Banksidia KlaCaLek (1921), Type species SiaUi immarginaiQ Say (misidentified as PerIa kamemis Banks), orig. de^ig., preoccupied (Ml]r, 1917). Banksiaf^a ClaASSen (1936). Replaoement for BanksieM KlapAlek. Caiaasoperiü Zhjltzüva(I967). Type specics Caui asoperSa sifinuiijera Zhilt^dva, monotypy, n. sy]i. Tytopyge KlapAlek (1913). Type species Tytopygepianidorsa KiapAlek, ori^. desig. ■A-duk fiataitus. - Tnocellatc, mediuiTi to J arge, pale yellow to black species. Wings pale to black, vcItis brown to black, Macropterous. Mesal occipital knob absent, Male genitalia.-T5 typically produced in to a rounded, excavated or cleft lobe; lobe absent in fcöJi.tfrfitJ, ipinulißra, and p\anidüTSa\ lobe or membrane of T5 with scnsilla basico-nka, T6-S typically scierotüed laterally and anteriorly, membranous mesally; T7-8 with at least rame manifestation of mesal sc I e rite covered with sensilla basiconica; T 8 mesal sclerite prolonged into a small posterior lobe. T9 typically membranous mesally with mesal sensilla basiconica patch. Hemitergites with short, broad tc slender, pointed anterior process, and well developed basal callus; apex o fan ten or process and basal eallus with sensilla basiconica. S3-7 with hair brushes. Penis envelope membranous; tube membranous with annulations, weakly to strongly sclerotized, or with distinct sclerites. Tube sometimes bearing small spiny lobes; sac lobed or sifftple, variously armed with [riangular spines or rounded, scale-like spines. Female genitalia. - Subgenital plate sJightly to strongly produced^ usually subtriangular and excavated, or notched apically. Vagina membranous, typically lined with fine golden seta-like spines; accessory glands present. Sperniathccal stalk shorter than vagina. Sperma-theca lisuaJly sausage-shaped, curled and bearing a few accessory glands. Egg. - Elongate oval, typically with stalked. Hanged collar, Collar typically short and wide, but long and narrow in media; rim typically irregularly incised. Chorionic surftce smooth, granular or with shallow to deeply impressed FCls. Micropyles usually large and numerous, set in posterior third; orifices sessile; canals usually not evident on surface. Anchor mushroom-like with short pedicel and globular cap. Larva, - Brown to dark brown with or without distinctive pattern. Occipital seta I row complete, Pronotum completely fringed laterally with stout spines. Abdominal tergal fringes of mixed long and short setae; terga with few sto tit intercalary setae; black clothing hairs present. Dorsum ofbody typically with mesal, longitudinal row of long, silky setae. S7 posterior fringe incomplete. Cercat swimming hairs present. Anal gills present or absent, Distribution. - Mainland Southeast Asia, .lapan, Taiwan, Soviet Far East, the Caucasus, Central Asia, eastern Nearctic. Valid species-iK Mintyla Wi; * chineitsis (Kt,APALEK}, n. comb, * ßavoiincta {McLArHLAN) * ßinwaa (Ba^ks) * hummetitii) (Naväs), n. comb, ^ prorsoieitix (Ma v As), radialis (Naväs), n. syn. * tchusa Stark Ä: Szczvtko * immarginaia (Say) indemaia Wu Claassin * japimica (Okamoto) Paragnetina Fig. 9 faragnesina rinaifle/init (McLachlan) J Japan]: Nymph habitus. I. Sivcc. B.P. Slark, S. Uchicia: Synopsis oflhe Worid Genera of Perlinae (Plf^ipltra: Pedidae) 23 * ardfljm.tfj (banüs) * tacrimosa (KLAfALtKj * ledoenm Stark & Szczytko * media (WALKtk.) * oi hrocephala KLAPALEK=6ttrmfln£T NavAs, e.fqwroU NavAs, oiiveri NavAs. mbriceps NavAs, n, syn. * pieli (NwAi) * ptanidorsa (KlahAlek) * spinulifcra (ZhiltzOva), n, comb. * iUfuAiV (OKAMDTOI * tinctipennis (McLalHlAN) Discussion. - Types off, acutistyla, F. intieniata, P. [rieli, P. lutescens NavAs, and P sc fienUingi ülapAlek are lost or unavailable. P. pieli is associated with our specimens on Lhe basis of a distinctive color pattern and figures of esternal genitalia in Wu [1938). in-i/entata may be synonymous with P. kiimmelina but until better material is available, this association cannot be formalized. P. acutistyla was recently described by Wu (1973) and the type shpuld stili ex.ist, bui has not been studied by us, The remaining names, P. iateyiens and P. srhetikiingi, are regarded as nomina dubia, P. io/haff (KlapAlek) is a member oft he»/^c-rofiL'uria« Jouktii complex, and eight additional nominal Paragnetina have been removed to Togoperla or Kamimuria in this study. They are: chekiangensh Chu, f/w^yara Wu & Claas-jormoxana ki.apaltk,/or/hnajf navas, infiimata navas, imegra klapalek, excavala KlapAlek, tonkinemis NavAs. The matter of fixing ü type for Bunksiana has been referred to the International Com mission on Zoological NDmendature by Stark & ZwicK, and the Tylopygtf synonymy resulted from a recent study by Zwic k (1988), 9.4. Oyuuiia KLAP,Ä!,ii:K Perta (Oyafniai iclapaltk (1907). Type ipecies Perla gibba Klapalek= Oyamia tu)fuhni (Mc Lac H lan), orig, desig. .'\dult habitus, - Triocellate, large, dark brown (Korean species with brightly handed legs, Japanese species with brown legs). Wings suffused, veins brown. Macropterous. Mesal occipital knob absent. Male genitalia, - T5 with a massive, bilobed, heavily sclcrolized process. T6 with or without a broadly biJobed process. T6-8 with mesa! membranous field, anterior and lateral margins sclerotized. Hemitergites massive, complexly bilobed and heavily sclerotized; apex of lower lobe and inner basa i margins of upper lobe with sensilla basiconica; lateral base of hemitergite with a sclerotized knob, S4-S wilh hairbrushes. Penis envelope with meso lateral patches of fine seta-like spines; tube armed with fine transparent spines; sac membranous, without armature; entire penis simple, without complcx lobes or sderites. Ferna I e gen i ta I ia." Subgen i (al plate s h ort, e üt ending o ver abou t a t h i rd of S9; ap ica I m argin rounded or slightly excavated. Vagina membranous with paired basal sderites, accessory glands present. Sperm a thecal stalk slender, shorter than vagina; spermatheca about equal to stalk in length, sausage-shaped, and attenuated apically; multiple accessory glands present. Egg. - Elongate oval with short wide collar, R.im slightly flanged, smooth and irregularly scalloped; ribs on collar numerous. Chorion mostly smooth or with irregularpunctations and irregular FC Is around collar. Micropyles subequatorial; orifices sessile, round and well defined; canals straight. Anchor mashroom-like; pedicel short; rap fiat, unspecialized. Larva, - Distinctly patterned in yelJow and brown; most abdominal terga with pale spots or transverse bars. Occipital setai row complete- Pronotal fringe of short thick setae, complete laterally. Abdominal tergal fringes primarily with short thick setae interspersed with a few long ones. Black clothing hairs present. S7 posterior fringe complete, Cereal swimming hairs absent. Anal gills present, DistribiitiDn. - Japan, Korea, northeastern China and Soviet Far East. Oyamia Fig. 10. Oyümia Itiguhris (McLachlan) (Japan]- Abdominal tip of female (a), vagina (b), abdomina! lip of rtia!e (c, in iatcraJ view d}, evened penis {e,f), and head and protEiora^ (b). J, Si tec, BP, Slark. S. Uchida: S^^lops^s of the World Genera oTPerlinae (Plecoplera; Perlidac) 25 Oyamia Fig. 1 U Oyamh !uf;iibri\ (Mc Lach LAN) [Japan.]: Nymphs habitus, Valid species.* /iigubrh McLfl.CKTAN ' Bamss = coream (Okamoto), ammca KlAPAlek, u, sjh, * seminigra (KiAl^ALEfc) = testryi (Nav\S), n. Syti. Discussion. - O. coreana and O. amtirica are synonymized with O, nigribasis. The type of cormna is very possibly lost It is not located at Hokkaido Univcvsity, Sapporo, where all survived speciti^ens of Okamoto'j stoneflies are boused. But his original description agrees well with nigriba-us in color patterns and male eenitalia. The type of amnrka also could not be examined by us. Tht synonymy is based on the agreement between the type of fif^ifta.vi.v and the rcdcicription of umiirica by Zhiltzova & ZAPEKlNA-dulkett (19S6). The synonymy oUezeyi with seminigra was proved by charBcteristics of the e^ (Isobe, 1988) frnm the leseyi holotype fern ale . 9.S. Aj^rietSna KlamaLEK Peria ^AgKeiinaJ KlapAlck (liJOT), Type speties Pi'da fleganiitlu KlapAlük, desig. KiapALek (1923), Phasgänop/iora Ki.apAlek (19 !4). Type speoiö Perla capitata Pictet, desig, KlapAlek Les TAGE (1921), Unnecessary replacement for Phaiganopkara KlapALek, not Pha:i- gonuphora G KAY, tfarrisiola Banks (1948). Type species Peria ßavescens Wals^t, orig. tlesig- Adult habitus. -- TrioceÜate, medium si^c, generally dark brown to black; sometimes with brightly banded femora. Wings dark, veins dark brown to black. Females macropterous, males macropterous or brae hypte reus. Mesa I occipital knob absent. Male genitalia. - Posterior margin T5 produced into paired low knobs, or a rounded, or shallowly notchcd lobe, T6 usually with broadly bilobed anterior sderite; T7-S sclerotized laterally and anteriorly, membranous mesally; T9 with slender mesal sclerite. Membrane of T 7-9 with or without sensilla basiconica; lobes of T5-6 with s ens i IIa basiconica, Hemiter-gites massive, usually extending beyond T9, sometimes compSexly lobed, and with large, spine-1 ike sensilla basiconica on apex, and scattered along dorsal margin; basal callus absent. S5-7 typically with ventral bruübes, usually strongest on S6. Penis envelope membranous; tube membranous but. armed mesally with paired patches of Ion g slender t ran s pa rent spines, and apically with small triangular spines; sac short, membranous and without armature. Basal sclerite present. Female genitalia. - Subgenital plate usually projecting over about one third to one half of S9; margin entire or eitcavated and frequently darkly pigmented. Vagina sometimes with paired basal or lateral scleritcs; accessory glands at anterior angles, Spermatheeal stalk slender and shorter than vagina; sperma theca slender ba.sa Ii y, enlarged and rounded apical i y; accessory glands present. Egg. - Elongate ovaJ, with butt on-I ike attachment of anchor; anchor large, umbrellalike. ,Anterior pole subtended by an inconspicous chorionic fold. Chorion granular and without FC Is. Mioropyles in posterior third; orifices small, sessile; canals s] anted. Larva. - Distinctly patterned in yellow and brown. Abdominal lerjja with posterior yellow bands or paired spots. Occiput with complete setal row. Pro not um completely fringed with mixed long and short setae. Abdominal tergal fringes with several long setae mixed with short thick ones. Black clothing hairs present. S7 posterior fringe complete. Cereal swimminjj hairs absent. Anal gilh present. Distribution, - Ncarctic, Palearctic, mainland Southeast Asia, Taiwan, ]. Sivec, B.P. Stark, S. Uchida: Synopsii oflhe World Centra of Periinae (Plecoptera; PerJidac) 27 Valid species," aculipenttii Kj.ApAe.RK * aeijuulis (Banks), n. tomb, * anttuf/pfiv ^H^GF.N) * artnata 1 Banks) * hrevipeitnii (NavAs) * capiiata (PictPt) ikryamJei (lM/\vAs) ' cifcumjcripui (KlapAlekX n. comb. Jubia WICK dupiLilyla fWu) * elegdiiiala [Kli^?Ai,EK) * exiretTia (NavAs) = fenilis (K1-ApAlek). cosUlata (NavAsJ. undata (KlapAlek). pedaia (Koponen), ? n. syn '-Q Agnetina Fig, 12- Agfietifiai'apilala (PicteT) [Canada]: Abdominal tip of female (a), vagina (b), head and firothora* (f)< abdominal lip of male (c, in laTerai view d); Agnetina brempefinis (NavAs) [Mongolia]: everted penis (el Agnetina Fig, II Agm^tintJ Jlavvxvrtf (WalshI [USA]: Nymph habitus. L Si>BC, B.P. Stark. S. Uchida; Synopsis of the World Gencin of PeiHinae (PIccoptera: Perlidat) 29 * ßaveitL'tis (Walsh) !^ladk(a (Wu) muhispinoaa (Wu) naraji {Wul, n, comb, * !)raeUi!a (klapalhk), n. comb. * cimJmttbenulaiu (WuJ n. comb-^pinaia (WLT) * MCfJie'" (KEMPIW) Discussion. - Stark (1986) noted the Nearctic species of Agneiina are apparently the siättr group of^, etegantuta ftom Eiirope. Earlier, Stark & Ga^urn {1976) suggested this might be due to a trans-Atlantic dispersal, but it now seems more probable that an Oriental, e/fgaw/u/ff-iike ancestor dispersed westward into eastern Europe and eastward across the Bering Bridge as pan of ths Pliocene fgunal e^tchange between Asia and eastfiiti North America. ZwiCK (1984) and Stark (1936) have recently reviewed European and Nearctic Agnclina. 9.6. Marihamea Ke.apAlfk Perh {Manhamea) KlapAlek (I9fl7), tyfk species Peria vitripen^h Burmüibter, desie, KlapAlek (I923J. PcHa (Ler/iaj Navas (1909). Tj-pe specics Urpa hcraiidi Maväs. AdüJt habitus, - Triocellate. medium size and generally dark brown lo black, Wings transparent, veins dark, costal field pale. Usually macropterous, some males brachyptcrous. Mesal occipital knob absent. Male genitalia, - Posterior margin T5 produced into a rounded or shallowiy notehed lobe covered posteriorly with sensilla bas iconics. T6-S sclerotized laterally and anteriorly, membranous mesally, dense mesal patch of sen sil la basiconica present on at least T7. T9 completely scicrotiicd around anterior margin. Hcraitergites bilobed in lateral aspect; ventral lobe slender, dorsal lobe broad; both lob« with apical sensilla basiconica. Dorsal aspect of heraitergites with inner basal callus coveted with sensilla basiconica. S6-7 with mesal hair brushes but less well developed on S7, Penis envelope membranous, tube weakly sclerotized, transversely annulate and unlobed; sac larger than tube and armed with variable size, thick triangular spines; basal sclerite present. Female genitalia. - Subgenital plate slightly projecting over anterior margin S9. Vagina membranous, slender accessory glands at anterior angles, S perma thecal stalk slender; sper-matheca small, apex rounded, accessory glands present- Egg, - Elongate ova!, with but t on-like attachment of anchor; anchor large, umbrellalike. Anterior pole subtended by chorionic fold. Chorion granular without FC Is, Micropyles in posterior third; orifices small with slightly raised lip; canals slanted. Larva, - Distinctly patterned in yellow and brown. Head mostly yellow, but dark between posterior ocelli. Abdominal terga with posterior yellow bands or paired spots. Occiput with complete setal rcjw. Pronotum completely fringed with mixed long and ^hort setae. Abdominal tergal fringes of long and short setae. Black clothing hairs scattered over dorsai body surface. S7 with posterior fringe interrupted. Cereal swimming hairs absent. Anal gills present or absent, Distribution. - Europe and Middle East. VaJid species- * beraudi (Navas) * jsJyiiV (Piltet) * fiitipenfiK (ßuRMEiSTER) Discussion. - Marthamea was redefined and the Palearctic species revised b> ZwiCK (19S4), who also noted Ihe great similarity in egg and larval characters with Agnetim, the apparent sisler group. Our study of t he types of Asian peri ids sometimes included in .Marthamea, indicates that all iDrientaJ species belong to other genera and that Marthamea is a European-Middle Eastern genus, Mariham^u is a pDtamophilic group, and according to Zwicii (1984) many European populations are endangered, or already extinct. Marthamea pig. 14 .Wart/wmra jf/ysi (Pectet) [Spainl: Abdottiitial tip of female (a), vagina (b), head and proihorax (c), abdominal tip of male (d), hemitergite in dorsal (e), and lateral vieiv (f), and everted penis I, Siv«. B.P. Stark. S. Uchida: Synopsis of the WoHd Gen eni of Perlin ae fPIecopteia; PeHidae) 31 Marthamea Fig, 15. Kiarihamea srp^-sii (Pictet) tSpain]: Nymph habitus. 9.7. Kamimitria Ki.AfALtK Perfa fKamimiiria) KlapAlek (1907). Type species Ferb libialii Pictet, desig. KlapALEK (1923). Adull habitus. - Triocellate, medium to large, pale brown to brown. Wings pale or suffused, veins brown. Females tnacropierous, maiejbrachyptetousar mai^ropterous. Mesal occipital knob absent. Male genitalia. - T5-8 unmodified; T7-8 sometimes with sensilla basiconica. T9 fre-quentlv with small mesa I lobe and median sclerite covered with sensilla basiconica. Hemi-tergites short, simple, finger-like, and without basal callus; apex with sensilla basiconica. S4-7 and metastemum with hair brushes. Penis envelope frequently with fine spines; tube sometimes with small paired sclerites, more typically membranous: tube typically armed with stout and small spines; sac shorter than tube, usually with a few simple lobes and variously armed; sac armature in some species massive, in others minute. Female genitalia. - Subgenital plate weakly to moderately produced, seldom extending over basal third of S9. Vagina membranous, accessory glands absent, Sperma thecal stalk small, slender; spermatheca a large inflated, bal loon-like structure. Egg. - Elongate oval with variable collar. Collar stalked or sessile, usually wide; rim usually flanged and weakly incised. Chorionic surface smooth to ornately sculptured; frequently sculpturing takes the form of large, shallow pits. Micropyles subequatorial; orifices usually sessile, canals straight. Larva. - General color brown with subtle pattern on head and thorax. Occipital setal row complete. Lateral pronotal fringe complete, consisting of mi.«d long and short setae. Abdominal tergal fringes variable; a few long setae on more basal segments; intercalary setae abundant, or sparse. S7 posterior fringe incomplete, Black clothirjg hairs present. Cereal stiim-ming hairs weakly to strongly expressed. Anal gills absent. Distribution, - Widespread over mainland Asia, Japan, Okinawa and Taiwan. Valid species.* amtiemi klapalek * brwineicarnis (KlapAlek), n, comb. * i oarciam KlapAlek fracva Harper * e.rö/j (McLachlan) = (uieicauiia KlapAleK., n. syn. * fiilwscens KlapAlek.^ /afjor Kla^Alek, n. syn, * formosana (KlapAlek), n. comb, (not Okamoto, 1913) himalayana Harpf.r * ii\fiimata (NavAs), n. Comb. * imegra fKiAPALEn.), n. comh, * kehtiUtnica KlacAlek * klapuJekia (Wu & Claassen). ti, comb. * kpiilii KlapAleK iiui Wl- = tm'vata Wu, spi'milata Wi;, n. syn. * lumlenta Zweck tiigrila Wl) onhvgofiia W;j * quaärata KlapAIEü sahlbergi Kotonen sentkosa Harper I, Sivec, B.P, Stark, S. Ucbidy; SjTiopsis of the World Genera oFPeHinae (PIccoplera; Ptrlidae) 33 * sH'kimensis (ENDtftLHiN), n. comb, * similis KlapAlek * tibialis (Pici Ei) irapezoidea Wu lubtrosa Wu Discussion. - The species lisl above excludes 14 Kammuria names proposed by Wu prior to 1962, for which type material is lost, and 2 Naväs sind 2 Klapälek Kamimifria under similar circumstances. We have also excluded 14 Asian spccies described by various authors Kamimuria Fig. Sfi. Kamimmia tibialis Peltet [Japan]: Alidorninal tip of female {a), vagina (b>, head and prothorax (g], abdomina! tip of male (c), in lateral view (d), and evened penis (e,f). Kamimuria F is. 17. Kamimiirici libiali.i PiCTET [Japan]: Nymph habitus. I. Si Vit, ri.K Slark, S, Uchida! Synopsis of the WoHd Gtnerj d Perlinae (pletnptcra: Perlidae) 35 in the late I91h or early 20tii century in the genus Ferla. Lost type material anJ inadequate descriptions preclude their inclusion in Kcimtfnvria or other perline genera at present. Peria nirvana Wanks,/', melanopkaltna N^vAs^and Kamitnuriajlavara NavAs, 1933 (not Perla IKaeTiimiiriaJßavata NavAs, 1923) are also excluded from Periinae and are considered as undetermined Acroneuriinae. Six other specics (Perla anamensts BamkS, P. shesloperüM NA-vAs, P. Slier tea IMavAs, Parafjnetina excavaia KlapAlek, Kamimnrta pola in i na NavAs, A'. ranjOia NavAs) are l^nown from female typtt which couid not be associated wiili any generic name. Our preliminary study of types, and other Kamimuria specimens, indicate several, rather distinctive, species groups are included. We presently recognize 7 groups; the lar^^e^t group ihiUiknla) includes 4 named species from the Himalayas, UcHlDA & ISOBEare revising the tihkiHx group from Japan. 9.«. Etrocorema klapäi.ek Etrocoremu Ki.apAlf.k (i'^o^t). Type species Etwcoretna ahenobarba Klapalek= Ochihopetma niffry- g/enkidaUl Monolypy. Euryptax KlapAlek (!909). Tvpe species Euryptax ixhrosiomu KlapAlek, Preoccupied tsrimpson, 1S59I), JVeoti!(n'/j/ii.vclaassen(1936). Replacemeni far Eurypiux Kiap vlek. Adult habitus. - Biocellate, medium to large size and generally brown to dark brown. Wings suftused brown with dark brown veins. Females and maies macropterous, Mesal occipital knot) present. Male genitalia, -T5-8 unmodifieil; T7-8 typically with patches of sensiila basiconica. T9 anterior scleritc produced mesally, partially dividing membianous field: usually a few sensilla basiconica scattered in membrane. Hemiter^ites simple, finger-like, extending to posterior margin of T9; apex with dense patch ofssnsilla basiconka. S6 and metastcrnum with liair brushes, Penis envelope membranous; tube weakly sderotized and trai3aversely annulate mesally; sac shorter than tube, apex expanded, multilobed and armed with thick spines; lar^st spines an tea pica!; fine long seta-like spines in lateral patches near base of sac; broad basal sc!crite present. Female genitalia. - Subgenital plate massive, completely covering S9 and usually extending beyond laieraf niar^jins of abdomen. Apex of plate with shallow, narrow notch. Vagina membranous with accessory glands: spermathecal stalk short; sperma t heca sniaäl, bulbous, Egg. - Elongate oval, with wide short collar, Rim of collar smooth with irregular incisions; flange absent; sides with numerous ribs. Chorionic surface smooth except for posterior poie which has shallow FCls surrounding clusters ofpits. Mieropylei subequatoriai; orifices sessile; canals straight. Anchor pedicel short, cap complexly and finely divided, appearing to consist of numerous fine strands; margins of cap with long finger-like projectionü. Larva. - .Mmost uniformly brown wiih subtle pattern on dorsal surface. Occipital ridge devoid of setae except near postocular fringe, Pronotal fringe complete, consisting of uniformly long, tl^ick setae. Abdominal tergal fringes of mi;(ed long and short thick, setae. S7 posterior fringe incomplete. Clothing hairs red-brown. Ana! gills absent. Cereal swimming hair? sparse. distribution. - Southea.st Asian mainland, Sumatra, Borneo. Valid species.-hoi hii (wii) • ftip'o^xnicuhium (Enoeriein) Etrocorema Fig. IS. ElrvCarema nigrosenkiilatum (EnderLEIn) [India]' Abdominal tip of" female (a), vagina (b), head and prothoras (c), abdominal tip of male (d), hemiiei^te (e), evened penis and ventral view of evened sac {g). Discussion, - ZwicK (1973) described the lectotype male of£, niswjienkithtum and in other papers (19S2b, d J 984) he clarified tKe systematics of t he genus placing several species and generic names in synonymy. The present concept of a single variable Etrocorema species throughout Southeast Asia should be tested by SEM examination of eggs from several populations when additional specimens are available. We observe rather conspicuous differences in head and leg colouration, and a few subtle differences in egg chorion detail and penis setation among the specimens; available to us. 1. Sivec, B,P. Stark, 5, Uchida: Synopsis of the Wodd Genera of Perlinac (Plecoptera: Perlidac) 37 Etrocorema Fig- ['S, Etrocorema nigrogeniivlarum (Enderlefn) [Thailand]: N^niph habitus, 9,9, Tetropina KjapAllk Teiropina KlapAllk (19093. Type species Ochthepeiina fitlgeicena Enderlein, orig. desig. Adult habitus. - flioceJlatc, medium to (arge, and genera;Jy pale brown; head and pro-notum usually with conspicuous pattern of dark on light background. Mesa I occipital knob present. Tentorial callosities adjacent to ocelli, directed posteriorly and longer than wide. Male genitalia. - T5-9 unmodified. T9 with sparse posteromesal patch of sensilla ba-siconica. He m it elites simple, short, rounded apically and densely covered with sensilla ba-siconica on apex. S6-S and mctastemum with hair brushes. Penis envelope membranous; tube armed with wide, rounded scale-like spines; sac shorter than tube, with large basal lobe; sac armed with variably sized spines in a continuous apical band; smaller spines nearer apex; basal sclerite present. Female genitalia. - Subgcnttal plate unproduced, or slightly produced. Vagina small, membranous without accessory glands. Sperm a thecal stalk longer than vagina; sperma t heca bulbous ätid Without accessory glands. Tetropina Fig. 20, Tesmpitia larvata Klapalek [Malaysia]: Abdominal tip of female (al. vagina (b), head and prothorax (f), abdominal tip of mate (c), herailcryte (d), and everted penis (e), i. Sivec, B.P. Siark, S. Uchida: Synopsis of iht: World Genera of Perlinat (Ptecopteii; Ptrlidae} 39 Tetropina Fig. 2J, Tetropimi hrxata ICi apAlek [Malaysia]: Nymph hahhus. Egg. - Elongate oval, with short, wide or sessile collar. Surface covered with shallow FC Is which surround shallow pits or raised processes. M icropyles in posterior third; orifices sessile, sometimes surrounded by FCI rosette; canals straight. Edosion Jine presenbelow micropy-lar row. Anchor cap complexly lobed; pedicel short. Larva. - Distinctly patterned in yellow and brown. Occipital seta! row complete. Lateral pronolal fringe incomplete. Abdominal tergal fringes of stout setae; stout intercalary setae mainly in anterolateral patches. Black clothing hairs present. S7 posterior fringe complete. Cerci without swimming hairs. Mesal longitudinal row of long silky setae on body dorsum. Anal gills present. Distribution. - Borneo, Malaysia. Valid species.* fuijfeaenii (En der lein) * kraepelini (KlapAlek), n. comb. * iäfviita KlapALEK Dis>*;ussion. - ZwiCK (1984) gave comparative figures for male genitalia and eggs of the Bornean species, T. larvata and Juli^cscens. T. kraepelini is tentatively placed in this genus on the basis of the overwhelming similarity of the egg to those of the two Bornean species. 9.10. ^eoperlops Banks Ncopcrhps BaNJ(S (1939J, Type species Neoperhps gressitü BaKJai;subequal to tube, apex (imied Hith a pair of lateraJ patches of slender spines. Baival sclerite absent. Female genitalia. - Subgenital plate unproduced. Vagina membranous, accessory glands absent. Sperm at hecal stalk longer than vagina and slender; sperm at heca small. Egg. - Elongate oval with short, wide collar. Collar rim flanged and shallowly incised, sides ribbed. Posterior pole deeply pitied; shallow mdistinct PC Is covcr most of surface. Mi-cropyles in posterior third; onflces sessile, canals straight. Anchor mush room-like; pedicel short, cap flat. Larva. - Distinctly patterned in yellow and brown. Abdomen with pale lateral bands extending to at least TS. Occiput with complete setal row. Pronotal fringe complete- Abdominal tergal fringes of uniformly short, thick setae. Black clothing hairs absent. S7 posterior fringe complete. Cereal swimming hairs absent. Anal gills present. Lateral submental lobe with basal suture- Distribution. - Circum-Mediterranean. VaEid spccies.-* cjchrarea (Kolbe) 9.13. Dinocras Kl.AP\I,tK Pcrla (Dirtocrcus) Kij^fAlek [1*^07). Type spBcics Perla cephahfe.i CurTis, orig. desig. Aduit habitus. - Tri ücel I ate, large and general 1yd ark brown. Wings suffused with brown, veins dark brown. Females macropterous, males macroptcrous or brachypterous. Mesal occipital knob weak. Male genitalia. - T5-8 unmodified, Sclerotized anterior margin T9 projecting into mesa! field, partially dividing terjjum; dense posterpmesal patch of sens ill a basiconica broadly continuous; posterolateral angles of T9 membranous. Hemitergites short, wide, rounded apically and covered apically with sen sil la basiconica; projecting portion of hemitergites longer than wide; basal callus absent, S6-7 with slightly enlarged mesal setal groups, not organized into brushes; S 7-9 sometimes with mesal patches of sen s ill a basiconica. Penis envelope membranous; tube short, unsclerotiied and without lobes; süc subequal to tube, apex armed with incomplete band of short spines, Hasal scleritc absent. Fem ale genitalia.-Subgenital plate produced over at least ha If of S9, un notched. Vagina membranous, without accessory glands. S perma t heca i stalk longer than vagina; spermatheca an enlargetj sac. Egg. - Elongate oval with short, wide collar. Rim smooth to irregularly incised, without flange; sides ribbed, area between ribs pitied. Chorion with shallow, indistinct FCIs. Mi-cropyles in posterior third; orifices semi-triangular, lower lip projecting above chorionic surface; canals straight. Anchor m us h room-like; pedicel slender, length about et^ual to cap diameter; cap consisting of numerous fine strands. I. Sivec, B,P. Slark, S. Ucliida; SjinOpsis oflhe World Oetiera of Pcrlinae (plttwptera: Pertidae) 47 Larva. - Typically dark browji with paired pale spots on abdominaJ lerp. Paäe areas on pronotal disc form a complex pattern; preoccipital area with large oval pale spots adjacent to eyes, but most of area posterior toetdysial suture brown. Occiput with complete setal row. Pronotal fringe complete. Abdominal tergal fringes of unitbrmly short, thick setae. Black clothing hairs absent. S7 posterior fringe complete. Cereal swimming hairs absent. Anal gills present. Lateral submental lobes with basal suture-Distribution, - Europe. Valid species.* ceifhaioles (Curtis) • firrm (PECTtT) • mv^acephaia (KlapAlek) Dinocras Fig. 27. Dinocräfs ntenacephala (Klai'Alek.) [Yugoslavia]: Abdominal lip of female (a), vi^ina (b). abdominal tip of matt (c), hcmilcrgite (d), everted penis (e), and head and prothorax (f). Dinocras Fi^ 28. Dinocras megacephaia (XlapAlek) [Yugoslavia]; Nymph habitus. 1. Sivec, fl-P. Stark, S. Uchittt; S>TiopMS oftbc World Genera orPerlinac (Plecoptsra; Perlidat) 49 10, Tribus NEOPKRIJNI I fl. l. Furcapwla SI VEt, n. g« n. Type species. - Tyhpyge bifisnaia Wu= Furcapt'rla bijufcala here designated. Furcaperla Fig. 29, Fareaperta bifurvaia (Wu) [China]: Abdominaj tip of fcmale (a), vagina (b), e^g (c), abdominal läp or male (d), hcmitergite {e}, everted penis (O, dorsal view of everted sac (s), and head and ptothorax (h). Adult habitus. - Triocellate, medium to large brown species. Wings suffused with brown, veins dark brown. Macroplerous. Male basal cencal segmerl. slightly enlarged. Mesa I oc-dpital knob absent. Male genitalia. - T 5,6 unmodified. T7 with a sparse mesa I patch of sens i 11a basiconica. TS with a sclerotized mesa I knob covered with sen sit I a basiconica. T9 with a widt mesa I sclerite, covered with scnstlla basiconica, and produced into low lateral knobs, Hemitergitcs with a pair of small finger-like projections. S4-7 and metaslernum with hair brushes. Penis envelope membranous; tube weakly sclerotiied, weakly annulate and with a small mesal lobe; sac shorter than tube, and bearing a pair of basal membranous lobes and 3 large ctaw-likescleroti7.ed,apical hooks; sac armed in basal halfwith triangular spines; basal sclerite present, Female genitalia. - Subgenital plate slightly produced into a short, arcuate, apical Iy notched structure. Vagina membranous with numerous anterior folds; accessory glands absent, Sperm at hecal stalk short; sperm atheca slender with a few long a[;cessory glands. Egg. - Elongate oval with slightly stalked, wide collar. Rim flanged, margin scalloped. Chorionic surface finely pLinctaie throughout. Micrupyles in posterior third; orifices sessile; canals straight. Larva. - Unknown. Distribution. — China, Valid species.* hifunaia (Wu), n. comb. DiscussioD.- This aberrant genus displays some Neoperlini characters, such as the scleroti^ed penis tube and the hemitergite form, but it also seems to be allied to the ^'eoper-hps-Tetropitiu group b> the metasternal hair brushes, The types of T btfuicaia are lost, but a series of 14 males and 12 females in the Museum Alexander Koenig iti Bonn, are consistent in hemitergal detail with Wu's (1947^8) description and figure, These specimens were collected by J, Klapperich from ftuatun, Fukien, China, from May 10, to June 5, 1937. An ad-ditiona! mak in the British M Lseum of Natural History was col lected by M. S- Young in Foo-chow, China, 1935-36. 10.2. IWeoperla NEEÖHAM P^eudoperta Banks (! 892). Type species Pcria mcipitalh Pictft, orig. desig. Preoccupied (Bronn, I ßAS; Pect et, 1845). Neopcrla ntedha?^ U90S). Repkcement for Fi^udüperia Banks. Ochihoperina Endühlüin (190?), Tit« species achfln>peiit)a (lei-iivnnii Empfri ein, orig. desig. Javanüa Kj_apAJ:Ek (1909), Type species Fcria caligala Biirmeistfr, monotypy. Tropidogympiax Tjpe species TTopidogynvplo-^ fusapes Em)e;ri-ein= O. aenpenni\ Enderle sn. Formtmm KlapAlek (19 [3). Type species N^operta haiakeyamae Okamoto, orig. dc^ig. Preoccupieti (spck sr, 1^111). Fwmtimä KlapAlek (1914). Replacement for Formosina Klaf>älek. Ocdi'ia Ki.APAlek (1921), Type species ^Jcoperla dolichocephttla KlaPALEK., orig. desig. Sirnpaperlü We; (1962). Type species Sitnpüperlu ithfctifa^ha Wl, monotypy, ti. syn. Adult habitus, - Biocellate, small to medium, usually pale brown, Exceptionally black. Wings suffused, veins brown, Maeropterous. Mesa! occipital knob absent, 1. Sivec, ß.P. Stark, S. üchick: SynnpüB of tht World Äncra of Perlinae (PIccopCera: PertidaeJ 51 Male genitalia, - T5-6 unmodtfied, T7 typically with projecting lobe or raised process. T8 usually with mesal raised process, varying from simple, slightly raised hump to elaborate, sclcrotised, tongue-like and sometimes bifurcaie proccss. T9 typically with slightly raised hum ps. T7-9 processes, or membrane, with patches of sensilia basiconica. Hem itergites with anterior process simple, finger-like; apical margins of basal piece covcrcd with sen si IIa ba-sicontca. Sterna without hair brushes. Penis envelope membranous; tube typically scleroti zed, at least basally, sometimes armed with lateral patches of spines pr spiny lobes; lobes sometimes unarmed or sclerotized. Sac usually longer than tube, variably lobed and armed. Female genitalia, - Subgenital plate unproduccd or weakly produced. Vagina with or without sderites, sometimes lined with fine spines; accessory glands absent- Spermaihecal stalk lined with fine, golden brown spines, or unlined; frequently long and coiled, sometimes short and uncoiled. Spermatheca reduced t o small membranous bag for species with complex stalk, and somewhat enlarged for other species into a slender, !ong, membranous bag. "Shape, collar form and chorionic surface detail variable. Frequently striate or with ornate FC Ii and punctation^, occasionally smooth. Collar present or absent, stalked or sessile. Micropyles in posterior third; orifices sessile or with raised lip. Edosion line present or absent. Anchor variable. Neoperia Fig. 30, Ni-opcrlaßarexcmsCHu [China]; .'Abdominal tip of male (a. in Jateral view b). head and prothorax (c). and penis (d). Fig. 3 K jVffl/if r/a obiiqua B^\^KS [Philippines]: Abdominal tip of male (a), evened penis [b), vagiria [c). egg (d)- I. Sivcc, a. k Stark, S. Uchida; Synopsis of the World Genera of Perlinae (PJecoptCTa: PcrJidac) Neoperla Fig. 32. Neoperla angidaja (W/vlkhr) [Sri Lanka]L Nymph hübiius. Larva. - Generally brown with subtle pattern Occipital ridge bare. Pronotum usually witliout complete lateral fringe. Abdoniinal tergal fringes typically of short, thick setae, S7 poSfterior fringe coniplete or incomplete. Certi usually without swimming hairs, but present in at least one species. Anal gills present. Black clothing hairs typically present. Distribution. - Nearctic, Ethiopian, and widely distributed over Asian mainland and southeast to New Gtjinea. Valid species. ~ One hundred eighty two Neoperla species are currently rccognized following wqrk over the past decade by Zwick, SrvEC, Si ark., and others. Included in this number are 19 species described by Wu, Chu, or Wsj & Claassen for which type material is lacking, and 29 African species names. ZwiCK is currently revising the African species and UCHIDA is revising the Japanese species. Much additional material from mainland Asia, particularly China, will be needed to clarify the situation in this region, SimpHperla is placed as a provisional synonym since the only known distinguishing character, absence of modified T7, is now known to occur in Neoperla iUk^i ZwtcK and tenuispina KlapAlek (ZwicK, 1983), 103. Chinoperfa Zvvjc k Smüfjerla Wu (1^48). Type species Sinoperh mgroßamta Wu, orig. dcsig. Preoccupied (PiNC, 192S), ChinopL'rh /wjcK (I'JSO), Replacement hr Sinopt'ria Wu- Adult habitus. - Biocellatc, small, brown to dark brown. Wings sutTused, veins dark. Macropterous, Mesa I occipital knob present. Sc vein usually ending in basal third of fore-wing, Male genitalia. - T5-8 unmodilied, T9 with slender, bil obed, mesa I sclerite arising from anterior margin, Hemitergites with long slender anterior processes approaching T9 sclerite; inner margins of basal piece of hemitergite with patch ofseniiilla basiconica. S3-4, sometimes 5 with hair brushes. Penis em/elqpe membranous; tube partially sclerotizcd basally; sac sub-equal to tube; armed near apex, typically with short spines but in at least one species with a dense cluster of long setal-like spines. Basal sclerite present. Female genitalia, - Subgenita! plate variable, covering most of S9, or unproduced, usually notchcd. Vagina membranous, without accessory glands. Spermathecal stalk short; spermatheca inflated, bal loon-1 ike. Egg. - Extremely variable in shape, chorionic detail and collar structure according to Zwicii(1982). Larva. - Unknown. Di,stribution, - Southeast Asian mainland, Sumatra, Borneo, and India, Valid specles,- * (Banks) * n/j^mvp f (Banks) rtigfiißävala (Wu) * ni^ifroiis (Banks), n, comb. = JunomcKiila (Wu), n. syn. * rediiaa (gtljskes) Discussion. - ZwiCk (1982) and ZwiCK & Sjvec (1980) recently gave details I'or several Chi naperi a species. The type species, C. nigrqfluvata, unfortunately, is known only from Wus (1948) figures, and is the only described Chinoperta which apparently lacks the L Sivec, B.P. Stark, S. UctiidaL Synopsis of the World Gtnem ot Perlinae (Plecoptera; Perlidae) 55 shortened Sr vein(ZwicK, 1982), We are provisionally placing 5" /«fictimatu/tj (Wu, 1973) as a synonym of C. nigrifi-ons (BA^fKS, 1939). Both species are from Hainan, and iherc is little to distinguish Wu's figure ofcJitcrnal male genitalia from ihc type of C, nigrifron^. Chinoperia Fig. 33. Chirmperia wfj^rrtt'/ij (Banks) [Imlia]: Abdominal tip of female {a}, egg (h), abdominal tip of male id), head and prijihoran (c), vcnirjl view ofpttiis (cJ, everted penis (f), fore wing (g); C. nimfron.'mph habiius. I. Sivcc, ft. P. Suirt, S Uchida; Sjnofteis of the World Genera Of Psrlinae (Plecopteni; PeHidae) 59 Egg. - Elongate oval with variable collar. Chorion smooth to ornate, Micropyles in posterior third; orificcs sessile. Anchor tap typically with tinger-like p roj ei; Lions; pedicel of variable length. Larva. - L) sually pale brown without distinctive pattern. Occipital ridgp with or without setae. Pronotum fringed primarily with long slender setae. Abdominal tergal fringes; unifgrm; terga with dense red-brown clothing hairs, S7 posterior fringe complete. C ere i typically short, stout and without swimming hairs. Anal gills present; gills PSC 2, 3 reduced; one pair PSC3 gills. Thoracic sterna] plates with posterior fringes qfsetae. Lateral lobes ofsubmentum with basal suture- Distribution. - Southern Oriental I stands and mainland Southeast Asia, Indian subcontinent, Sri Lanka and the Himalayas. Valid species, - Thirty four species have recently been described by Zwtt K. (1982, 1983, 1986). ZwiCK &SivT;c(l985)andSTARK(l9S3>, ZwicK (19S2) recognized 7 species groups based on egg and penis characters, t T, Mimperla Kawai Xfiniperk Kawai ([967). Type species ^fimpe^h japanica Kawai, orig. desig. Adult habttus.-Triocellate, small and generally pale with dark markings. Wings transparent, veins except costal area brown; C and Se yellow. Macropterous, Male Benitalia,-T3 with a slightly sclerotized, weakly produced mesa I lobe, T4-8 with sclerotized anterior mai^in usually broadly bilobed; mesa I field of T4-S membranous, T 9 with mesa I membranous area bounded by raised sclerotized bars. Hemitergites slender, finger-1 ike, extending forward over posterior margin T9. S3-4 with mesa I hair brushes. Penis characters unknown. Female genitalia.-Subgcnital plate truncate, short, and barely projecting over anterior margin of S9. Vaginal characters unknown. E!^.-Lin known. Larva,-Unknown. Dist r ibu tion ,-i a pa n. Valid spccicš,-iapmKu Kaw.m Discussion,- This small, distinctive species is known only from Kaw.m's (1967) description ofa series from the now polluted Lake Biwa outHow. Speciemens have not been available for our study. 12. Species not included in the lists of valid specie* The following iist includes species unrecognizable from the criminal deseripiions for which the typic material is losi: A^eiina ckrysodes NaVAS. 1919; Pamgnetifia hitexfns NavAs. 1911;/'. Ki.ai'Ai,eti,1921; P. tmikinensis NavAs, \9\')\Pi'ila aegyptiaca PiCTET, iS4l; P. upkala NavAs, i^J fS; j^. icidavetam Mc-Lachlan, \ ü.ly. p. i-arietom Banks. p. caucafica GutRr^, 1S38; p eaudma Ki ap^lek, nlhilala NavAs. I9]9; f. tocandka McLaLüLAN, 1875; P. iowf.ftoJt/ wu, 1937; F. coaam Nav«. I91S-. cymbek Neeuiiam, 1909; P.Jlavam NavAs, l?>23 {not Kcimimuna^cii'hm NavAs, 1933); A im-mtrsa mtlah^ilak. i87s:p, mipuMata Pfc-nrr, t!t41; p. infiimala navas, 1923; P. aitw meedh;\w. iw; p. Claassen. 1923; p. picKii Lltas, 1849; p. isenitlei Pictet l«4i; f. .iiuna^iva samal.|935; P. simphxCnVt 1929; P. .vetmcta banks, 1914; Phasgaiiophora media Wu, 1935; Kamimuria aJraurnü KLA?Altk, 1912; A", t fa-m Wu, 1947; K. ckin^echenmnLiWu, !93S; K. ihuiigftämhatia W[J. 1938; A". croisispina Wu, 1947;A',yraMnei[ Wu, 1935",K, kiriwmm Wu, 1938; fT, hmgispina 1947; K. magna Wu 1938; K. manchiiriana Wu, I93K; K. melli Wu, 1935; mi^riceps NaVaS. I?i9; A'. riolalJs NavAs. 1927; a: roperh üf Sumiiira and Java (Indonesia) (PIecopierü: Perlidae), Spi-xiana. 6: 167-204. ZwrcK, P., 1 Manhameci ijeraudi (^livas] and its european congeners (Piecoptera: Per-Hdae). Annal. Umnol., 20: 129-139. ZwicK, P., 1984b: Notes on the genus Agnetina (= Ffiaxganophora) {Plecoptera' Perlidae), Aquat. Insects, 7U79. Zw I PK, P., 1984c; The genera Tctrppina and Neoperhpi (Plecoptera; Perlidae). Aquat. In-sccts,6: 169-176. ZwiCK, p., 1985: The Borneati species of the stonefly genus A'ew/jcr/c (Piecoptera; Perlidae). Aquat. Insects, 8: 1-53. ZwiCK, P., 1986: Contribution to the knowledge of Phanoperia Banks, 193B (Plecoptera: Perlidae). Mitt. Schwciz. Ent. Ges., i5J-158. ZwicK. P., in press: Notes on Plecoptera (16), Tyhpyge Klapälek; a synonym ofPara^^netina Klapälek. Aquat. Insects, 10. ZwicK, P., & I. Sivec, 1980: Beiträge zur Kenntnis der Plecoptera des Himalaja. Ent. Ba-siliensia,5: 59-13S. Zwict. P., & I. Sivec, 1985: Supplemente to the Perlidae (Pleeoptera) of Sumatra. Spixiana, 8; 123-133. k Fig. 37. SEM phoiomkrographs of eggs: Togoperh s.p. iThaüand]; Egg lateral. 200s< (I); Tofioptrla lim-hala (PiCTET) [Japan]; Egg lateral. 150<{2); Paragnfütia media Walkes [USA]: Eg« laterai, 250* (3); Oyaimu sp.: Egg lateral, 3B[lx (4), collar, lOOOx (5), detail ofthoiHon and micropyks, 2lKJ0x (6); Mar-ihamea fterdUi/f (NAvAsJ^LebanöTi]. P.ge lateral. 306x(7Xp'sia]; lateral, 2(Wk (10), collur, 4()0y {H)t Teiropina sp. [Malaj'sia]: Egg lateral, 200)( (12); Peria hunni'iiwnam Q.Ai^SSF,N ISwitaerlandl: Egg bleral iOÜx (13), collar end (14), delail of chorion and raicrop>leJj, 700f( (ISJ, Pi'tla paiUda Gl^rin [Yugoilavia]: Egg lateral. 230* (16), daail of chorion and micropyles. 95ÜX (tj^ Perk grandis Rambl K [SwitJcrland]: Egg lateraf 270x OS). '"m Fig. 39. SEM photomicrographs ofc-egi; Eoperh ochrm-ca (Külbe) [Algeria): Egg lateral. 220s<; 19}; ßi-11 tx-™ megacephata (Ki.apA.lek) ISwilzerland]: Egg lateral, 200x (10). collar end, 700it (J 1V. Neoper}a [CMna]; Egj kteral,27(b<(22),d«ail orcEiorion and micnopyle, ! 2IXlx(33), polar end. 550>!(34); Neoperla iahu Staak [Thailand]: Egg bteral, 300x(2S), collar ei>d, 750x(2ft>; Neoperla sp, [Sumatra]; Eggs lateral, {27}. ■ "f" ■ ____L'^__