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Russian Entomol. J. 16(3): 245-258

ї RUSSIAN EN TOMOLOGICAL J OUR NAL, 20 07

Review of Ameletidae (Ephemeroptera) of Russia and adjacent lands Обзор Ameletidae (Ephemeroptera) России и сопредельных стран Nikita J. Kluge Н.Ю. Клюге
D ep a r tme nt of Ento molog y, Sa int- Peter sbu r g Sta t e Uni ver si ty, U niver sitet sk a ya na b. , 7 /9 , Sa i nt-P eter s bu r g 1990 3 4 , Ru ssia . Ema il: k lu ge@FK13889 . spb.edu , - , .- , ., 7 /8 , .- 199 034 Ru ssia.

KEY WORDS: Ephemeroptera, Ameletidae, Metreletus, Ameletus, Metreletus micus comb.n., Ameletus altaicus sp. n., Ame letus montanus rossic us ssp. n., Ame letus montanus arle cchi no ssp.n. : Ephemeroptera, Ameletidae, Metreletus, Ameletus, Metreletus micus comb.n., Ameletus alt aicus sp. n., Ame letus montanus rossic us ssp. n., Ame letus montanus arle cchi no ssp.n. ABSTRACT . Revised generic diagnoses of Metreletus an d Ame letus are given. T he species, or igina lly described a s Ame letus mic us, i s tra nsfer red to genus Met relet us. Basing on all stages of developmen t, th ere are described a n ew species Ame letus alt aicus sp. n., as wel l as new subspecies A. montanus rossic us ssp. n. a nd A. montanus arlec chino ssp.n., which inhabit in Siberia and Russi an Far East . Brief cha racter istics of other species an d subspeci es, distr ibut ed on ter ritor y of Russia, are given. Characters of l arval geni tal buds as such, are used in taxa dia gnoses for the first ti me. . Met relet us Ame letus. , Ame letus mic us, Met relet us. A me le tus alt aicus sp. n., A. montanus rossi cus ssp.n. A. montanus arlec chino ssp. n., . Ame letus, . . recently revised by J. Zlot y an d G. Pritchard [Zloty, 1996; Zl oty & Pri tchar d, 1997]. Asian species were described from Japan [Imanishi, 1932, 1933], Taiwan [Kang & Yang, 1994] and the former USSR [Ba jkova, 1976; Brodsky, 1930; Kluge, 1979, 1982; Sinitshenkova, 1977, 1981; Sinitshenkova & Tshernova, 1976; Sinitshenkova & Varykhanova, 1989; Soldan, 1978; Ulmer, 1927]. The present paper con tains descript ions of new ta xa an d a review of formerly known taxa from Siberia and Russian Far East. All material examined is deposited in the Zoological Institute of Russian Academy of Sci ences, in Sai ntPet ersburg; t empor arily it l ocat es in the depar tment of ent omol ogy of Sa int Petersburg Universi ty. In th e lists of materia l examin ed, the following arbitra ry signs are used: Im -- imago; S -- subi mago, L -- larva, L-S -- subi mago rear ed fr om la rva (with lar val exuviae); S-Im -- imago rear ed fr om subima go (with subimaginal exuviae); L-S-Im -- imago reared from larva (with larval and subimaginal exuviae); L/S -- mature lar va wi th developed subimaginal cuticle. Family Ameletidae McCafferty, 1991
H iera r chica l na me: Ameletu s/fg1 ( incl. Me tr ele tus ) REFERENCES. Klu ge et al., 19 95 : 1 11 , Figs 1 8- 22 (egg, la rva , subima go, imago) ; Kluge, 20 04: 8 0, Figs 11, 20 , 21B-C ( la rva , su bima go, ima go).

Introduction
For the long time, mayflies, which are recently attributed to Ameletidae, wer e att ributed t o a family Siph lonur idae. In such wide sense, Siphlonurida e s.l . represen ted a plesiomorphon [about t his term -- see Kluge, 2004]. Because of this, recen tly such t axon is not recogniz ed, being split ted i nto sever al sm aller holophyletic fam ilies -- Siphlonuridae s.str ., Am eleti dae an d oth ers [Kluge et al., 1995; Kluge, 2004]. Ameletidae are distributed in Holarctic and Oriental Region, havi ng m ost n umber of speci es (about 30) in Nor th Am erica, less number of species in Asia and two species only in Europe. North Ameri can speci es were

La rva. Larva retains primary swim min g siphlonu roid specialization. H ead h ypogn athou s. Mou th apparatu s h as un ique specialization: labrum elongate; mandibles elongate perpendicu lar to their axis of articulation , w ith th in weak in cisor and kinetodontium, prosth eca on left mandible setiform, on right m andible lost; su perlinguae and hypopharynx elongate; maxilla elongate, w ith apical margin widened, pectinate setae of apical-ventral row elon gate an d form filtering apparatus, m axillary canin es com pletely lost, only one vestigial dentiseta retain ed. Maxillary palp retains 3 segments. Un like other mouthparts, labium n ot elon gate; labial palp retain s 3 segmen ts. Patella-tibial su ture retained on middle an d hind legs. C law s sligh tly curved. T ergalii lost ability of


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Nikita J. Kluge
contiguou s condition. Fore wing with anastomosed veins in pterostigma. Hin d w ing well-developed, as long as 0.35-0 .4 of fore w ing length (even in micus [Ameletus], wh ose fore wing lost trian gular shape). Tarsi 5-segm ented, 1st segment fu sed with tibia. All claw s ephem eropteroid. In male imago, styliger dorsally w ith a uniq ue mem branous area. Gon ostylu s retains 2 distal segments. Paracercu s vestigial. Comp osition and d istribu tion . D istribu tion of Am eletidae is lim ited by H olarctic an d O rien ta l R egion . Am eletidae are d ivided in to M etreletus an d A m eletus (see belo w ).

rh ythmical respiratory movemen ts. All th ree caudalii have su bequ al len gth, with dense prim ary swimming setae. Su bimago and imago. Meson otal su ture stretched backward and pointed medially. Both in imago and subimago, lateroparapsidal su tu re elongate; in subim ago, lateral pigmented area of m esonotu m inclu des entire su blateroscutum an d su bmedioscutum up to m edioparapsidal su ture (similar form of this area in Isonychiidae on ly). An terior paracoxal su ture complete. Both in imago an d su bim ago, epimeron of mesoth orax with a un iqu e membranous area between an epimeron and katepim eron. Furcasternal protuberances retain

Ta ble 1. Cha rac ters of sp ecie s of A meletus o f Ru ssian Fede ration , Ka zakhstan a nd Mon golia. Та бли ца 1 . Приз нак и ви дов Ame letus Р осси йск ой Феде рац ии, Каз ахс тана и Мон голи и.

Expla na tion for the T a ble 1: 1. Egg sha pe: "O" -- ellipsoid ( not fla ttened) (Fig. 17 ); " I" -- fla ttened, in a for m of ellipse-like biconvex lens (Figs 18 -1 9). 2. Ea ch polygonal cell on egg su rfa ce: "+ " -- with papilla; "-" -- withou t papilla (Fig. 17 ). 3. On egg surface round pa pilla-bea ring cells ( other tha n large papilla -bearing cells on one pole) : " +" -- prominent, with bor der s a nd papilla e distinctly higher tha n polygona l cell (Figs 17-1 9) ; " - " -- absent or not higher tha n polygona l cells. 4. On egg surface la rge pa pilla-bea ring cells loca ted on one pole: "+ " -- la rger than other round cells and higher tha n a ll cells (Figs 1 7-1 9) ; "- " -- absent or not la r ger than others. 5 . La br um: r atio of length to width. 6 . Ma ndible: "+" -- proxima lly with da rk ening or whole pr oximal pa rt dar kened; "- " -- pr oximal pa rt without cu ticu la r pigment except for a rea close to mola. 7 . Ma ndible: " +" -- with a rched da r k ening str etching from dista l-later a l ma r gin to mola , or a t lea st with da r k ening a dja cent to distal-la ter a l ma rgin; "- " -- dista l ha lf withou t cu ticula r pigment. 8. Middle dark ba nd on lar val femur : " +" -- femu r with a distinct dark ba nd or with compact da rk macula in middle; "-" -- femur unicolour or with long dar k macula. 9 . La rval a bdomina l terga: "+ " -- with spine-like setae; "- " -- without spine-like setae. 1 0. La rva l abdomina l sterna : "+" -- with spine-lik e seta e; "- " -- withou t spine-lik e setae. 11 . T ergalius II: "s" -- la rger than terga liu s I , but sma ller than ter galius II I a nd next ones (Fig. 6) ; " l" -- the longest, longer than tergalius Ш and other s ( Fig. 13 ). 12 . T ergalii II I-VI: "() " -- with anal rib just on a nal ma rgin; "())" -- with anal rib near a nal ma rgin ( Fig. 7 ); "(I)" -- with ana l r ib fa r fr om anal ma rgin ( Fig. 14 ). 13 . Subimagina l wings: " +" -- each cr ossvein bordered by dark ; " -" -- wings uniformly coloured. 14 . I magina l wings: "+" -- each crossvein bor dered by da rk; "- " -- wings uniformly coloured. 15 . La ter a l lobes of penis: " S" -- sigmoid, i.e. cu r ved dor sa lly, with a pices somewha t cu r ved ventr a lly; " (" -- distin ctly bent medially ( Fig. 11) ; "I " -- in ventra l view nea rly str aight or only slightly bent media lly (bu t bent dorsally) ( Fig. 16 ). 16 . Ventral plate of penis: "I" -- with la rge apica l stra ight spine; ")" -- with la rge apical ar ched spine; "-" -- without apica l spine: either without spines or denticles, or with sever al equa lly sma ll denticles ( see 1 7-1 8).


Review of Ameletidae of Russia and adjacent lands
L arval mou th ap pa ratu s. Mou th apparatu s, bein g m odified (see above), has rath er un iform structure in all Ameletidae. Labrum len gth is species-specific an d varies from 0 .65 of w idth (in N orth Am erican species A . m aju scu lus) to equ al to w idth (in A . p arvus, A. atra tus an d M etreletus). Man dibles (see above) have un iform sh ape in all species. Zloty [199 7] regarded relative len gth of denticles of incisor of left m andible as a species ch aracter: in A . celer and A. m aju scu lus secon d denticle is th e longest, w h ile in other N orth American species first denticle is th e lon gest; actu ally, in so m e of th e spe cie s exa m in ed b y m e (A . ce ler, A. a ltaicus, A . cam tscha ticus) th is ch aracter varies in dividually. Maxillae (see above) also h ave un iform sh ape in all s pec ies . M ax illar y pa lp is 3 -seg m e n t ed ; S in it s he n k ova [1981 ] regarded sh ape of apical segm en t to be speciesspecific, bu t th is ch aracter appeared to be in dividu al [Klu ge, 1 995 ]. Labiu m somew h at differs in M etreletus an d A meletus (see below ). Labial palps are 3 -segmen ted; Sin itsh en kova [198 1] regarded relative len gth of segm ents to be speciesspecific, bu t th is ch aracter appeared to be in dividu al [Klu ge, 1 99 5]. L a rva l cu ticu la r colou r pa ttern. As in othe r in sects, cu ticu lar colou r ch an ges du rin g m ou ltin g cycle (bein g ligh ter after m ou lt and darker before m ou lt); as in oth er m ayflies, larval cu ticu la r colou r pattern does n ot pa ss to th e n ext stages -- su bim ago an d im ago. U su ally the m ost part of larval cu ticle is pigm en ted; th is backgrou n d pigm en tation can be f ro m ligh t t o da rk br ow n , v ar yin g in d if fe rent species, in dividu als an d m ou ltin g ph ases of th e sam e in dividu al; h ere t h is pigm en tation is called d arken in g. Some areas of cu ticle lack pigm en tation , an d on liv in g larva th ese places are ligh t w h itish or yellow ish ; h ere th ey are called b la n ks.

247

An tenn a can h ave the follow ing colour pattern: scapus (wh ich is very sh ort) an d pedicellu s are partly darken ed; proxim alm ost and distalmost segments of flagellum are ligh t, wh ile middle segmen ts of flagellum are darker or partly darken ed. This pattern is expressed in those specimen s of Ameletus w hose cu ticle has con trastin g darkenings and blanks, but in specim ens with n on-contrasting cu ticle anten nae are un iformly coloured. Anterior surface of m an dible can have tw o darkened areas: proxim al darkenin g occupies most part of proxim al half of m andibles and adjoins its lateral margin; distal darkenin g is arched an d stretches from lateral margin of distal h alf of mandible tow ard mola. Most species have th e both darken ed areas, wh ich are often united into an in tegral darken ing; A. in opin atus has no proximal darken ing; A. mon tan us arlecch ino su bsp.n. an d A . m . ro ssicus sub sp .n. have no distal darkening (see Table 1 ). In most species of Ameletus, certain abdom inal terga have a pair of isolated roun dish blanks and an u npaired blank on backgroun d darkenin g (Fig. 14 ). As in other mayflies, each abdominal tergite can bear a pair of obliqu e submedian stripes diverging posteriorly; in the cases when they are darker th an th e backgrou nd darkening, they are visible as dark obliq ue stripes (Fig. 7 ). La rval colou r pa ttern of living tissues. U nlike some other mayfly larvae, larvae of Am eletidae h ave no marked hypodermal pigmen tation. Bu t in som e species abdom in al nerve gan glia are distinctly darkened and w ell-visible throu gh hypoderm and cu ticle as blackish spots. Like hypoderm al colour, this n erve colou r does not depend u pon m oulting cycle and passes from larva to subimago and imago. L a rva l te rga lii. In all Am eletu s/fg1 each tergaliu s III- V II h as a stron g costal rib on costal m argin an d a strong

17 . Ventral plate of penis: " +" -- with sever al sma ll denticles in pr oximal pa rt (Figs 11- 12, 15 -16 ); "-" -- withou t small denticles in pr ox ima l pa r t. 18 . Ventral plate of penis: " +" -- with sever al sma ll denticles in distal pa rt (Fig. 20) ; " -" -- without small denticles in distal par t. 1: 1. : O -- ( ) ( . 1 7); I -- , ( . 1 8 -1 9 ). 2. : "+ " -- ; "- " -- (. 17 ). 3 . ( ) : "+ " -- , , (. 17- 19) ; " -" -- . 4. , : "+ " -- ( Figs 17 -19 ); "- " -- . 5. : . 6. : "+" -- ; "-" -- , . 7. : "+" -- , - , , - ; "- " -- . 8 . : " +" -- ; "- " -- . 9. : "+" -- ; "- " -- . 10 . : "+" -- ; "-" -- . 1 1. II : "s" -- , I , , I II ( . 6 ); " l" -- , I II (. 13 ). 12 . II I-VI: "() " -- ; "())" -- (. 7 ); "(I)" -- (. 14 ). 13 . : "+ " -- ; "-" -- . 14 . : "+ " -- ; "-" -- . 15 . : "S" -- s- , .. , ; "(" -- ( . 11 ) ; " I" -- ( ) (. 1 6). 16. : "I" -- ; ")" -- ; "-" -- : , (. 17-18). 17 . : "+ " -- (. 11- 12, 15 -16 ); "- " -- . 18 . : " +" -- ( . 2 0); "- " -- .


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Nikita J. Kluge I. Genus Metreletus Demoulin, 1951
H i er a r chica l na me : Me tr e letu s/g( 1) REFERENCE. Kluge, 2 004 : 81 , Fig. 20 D (larva, ima go).

an al rib eith er at a distan ce from an al m ar gin , or on an al m ar gin (Figs 7 , 14 ). T ergalii I an d II are w e aker t h an oth ers, w ith co stal an d an al ribs vestigial or lackin g. In m o st s pe cies t er ga liu s I is t he s m a lles t, t er ga liu s II is larger, bu t m uch sm aller th an tergaliu s III an d n ext on es (Fig. 6 ). In tw o su bspecies -- A . inop ina tu s lab ia tus an d A. m on tanu s a rlecch ino sub sp .n. -- tergaliu s II is stron gly elon gated, bein g th e lon gest, bu t retain s its w eak con sisten ce (Fig. 13 ). Ma le g enitals. In im ago an d su bim ago, styliger has wide median in cision and a pair of widely diverging lateral projections (Figs 9, 16). In Metreletus, larval styliger has shape similar to imaginal one (Figs 1, 2). In all Ameletus larval styliger differs from imaginal on e -- its median in cision is mu ch narrower, an d lateral projections are less divergin g, more prominen t, pointed an d sometim es serrate (com pare Figs 8 an d 9 ). In imago and subimago, each gonostylu s h as two distal segm en ts (plesiom orph y amon g E ph em eroptera); in larva each gon ostylus has one in distin ctly separated distal segment, which corresponds to two distal segments of winged stages (compare Figs 8 an d 9 ). Im portan t spec ies ch aracters are foun d in pen is stru ctu re. Im agin al p en is bears a pair of sclerotiz ed lateral lobes located laterad -distad of gon opores, an d a pair of m em bran ou s or s cler ot ize d ven t ra l plat es lo ca te d pr ox im ad of gon opores [Zlot y, 1996 , 1997 ]; by sclerotiz ed pen ial arms th e pen is is ar ticu lated w ith corn ers of st yliger an d w ith IX abdom in al t ergite [Klu ge, 2004 ]. Su bim a gin al pen is is n on -sclerotized ; it h as lateral lobes of th e sam e size and sh ape as im agina l on es; as in oth er m ayflies, it lacks pen ial ar m s . L ar va l p en is is e ven ly sc le ro tiz ed , m uch sho rt er th an im agin al an d su bim agin al. Sh ape of larval pen is is m ore sim ple t h an in im agin al on e an d differs in M etreletus an d A m eletus (Figs 1-2 , 10 ). V en tral p lates in various species are eit h er sm ooth , or bear ch aracteristic den ticles. E ach su ch den t icle develops on a place w h ere su bim ago h as a sm all stou t spin e-like seta. T hu s, su bim agin al pen is bears spin e-lik e setae in th e sam e nu m ber an d arran gem ent as im agin al den ticles; th ese su bim agin al sp in e-like setae are h eavily sclerotized an d w ell-visible on backgroun d of th e re st su bim agin al pen is cu ticle, w h ich is n on -s clerotized. Larval p en is also can bear spin e-lik e setae, w h ich correspon d to t h e su bim agin al spin e-like se tae; nu m ber of th e spin e-like s etae on larval pen is is eith er th e sam e as in su bim ago, or less (com pare Figs 10 an d 1 1 , 20 an d 21 ). T h e presen ce o f com plete set of spin e-like setae in su bim agin al stage allow s to kn ow ou t arran gem en t of den ticles on im agin al pen is, stu dyin g su bim agoes or m atu re n ym phs w h ich h ave dev eloped su bim agin al cu ticle in side. O th er featu res of pen is stru ctu re, especially sh ape of its lateral lobes, are so s tron gly ch an ged du rin g tran s form ation from larva to su bim ago an d th an to im ago, th at exam in ation of larvae an d sub im agoes does n ot allow to d eterm in e th eir im agin al c on dition. E gg. Egg su rface has reticulation con sistin g of 2 kinds of cells -- polygon al and rou nd ones; inside each roun d cell locates a papilla (Fig. 1 8); bottom of polygonal cell can be eith er flat, or produ ced to a m ore or less prom in ent protuberance or papilla. Size of cells and papillae varies am ong species. Most species have eggs of a usu al ellipsoid sh ape (Fig. 1 7); in some species eggs are flat (Figs 18 -19 ) -- th ese are East Asian species A . m on tan us, A . ced ren sis an d A. p arvus, an d N orth American species A . celer, A . coo ki, A. spa rsa tus, A . suffu sus.

CH AR ACTE R IST IC S. T ill rec en tly, M e trel etus w as kn own as a single species ba lcan icus [Metretopus] and w as separated from Ameletus by th e followin g ch aracters: (1) In cu bital field of fore w ing, instead of vein s going from CuA to wing m argin, there are one or tw o in tercalaries (Fig. 4). (2) Larval claw h as a regular row of denticles. (3) Glossae are n ot truncate, apically-ventrally w ith num erous irregular pointed setae. Now I am placing here a second species -- micus [Ameletus]. It also has 2 in tercalary-like veins in cubital field of fore w ing (Fig. 4 ). But structure of claw s and glossae of M . m icus are u nkn own , becau se all legs and labium of the single kn own specim en (holotype) were detached and lost. So it is not qu ite clear, if the second an d the third characters really belon g to th e w hole genus Metreletus, or they occur on ly in M. ba lcan icus. In addition to the th ree ch aracters listed above, now we can add th e follow ing ch aracters com mon for the both species of M etreletus: (4 ) Imagin al pen is h as ven tral plates very lon g an d blun t (th is is reliably kn ow n for M . b alcan icus only); each lobe of larval penis h as a promin ent ven tral plate separate d by in cision from lateral lobe (Figs 1-2) (un like larval A meletus, w h ose ventral plates are not separated from lateral lobes -- Figs. 10 , 2 1). V entral plates of penis have n o den ticles or spin e-like setae (un like some A meletus). (5) Labru m h as a deep apical in cision (Fig. 3) (un like A meletus, w h ose labru m h as a sh allow apical in cision ). (6 ) Larval abdomin al cu ticle h as no blanks an d dark stripes: terga h ave on ly diffu se m edian darken in gs, sterna colou rle ss. CO MPOSITION AND DISTRIBUTION. Metreletus h as Trans-Palaearctic distribution an d in clu des two species: (1) West-European moun tain species ba lcanicus U lmer, 19 20 [Metretopus] (= hessei Fizaine, 19 31 [Ameletus] = go etgh ebu ergi Lestage, 19 38 [Metretopus] = hu ngaricus Ujhelyi, 19 60 [Metreletus]) and (2 ) Far-E astern species micus Bajkova, 19 76 [Ameletus].

I.1. Metreletus micus (Bajkova, 1976 ), comb.n. (Figs 2-5)
RE FE RE N C ES . Ba jk ov a , 1 9 7 6 , 5 8 3 , Fi gs 8 - 1 0 ( l a r va ); Kluge, 1 995 : 9 , Figs 4- 5 ( genita l buds). MATERIAL EXAMIN ED. RU SSIA: b asin o f river Ussur i, river Va k, 3.VIII.19 58 ( O. Bajkov a) -- 1 L/S# ( holoty pus).

La rva. All abdominal terga an d stern a w ithout den ticles on posterior margin s; posterolateral spin es absen t on all segmen ts (unlike all other Ameletidae, wh ich have posterior tergal denticles an d posterolateral spin es at least on posteriormost abdom inal terga). T ergalii (at least V-V I) very long an d slender, each longer than tw o segments, with len gth three times exceedin g width (un like all oth er kn own species of Am eletidae, w hose tergalii have length 1 .5-2 .5 times exceedin g width ); anal rib very slen der, separated by a small distance from an al margin (Fig. 5 ). Su bima go a nd imago (basing on w ings extracted from m ature larva). Fore w in g oval, w ithou t prominen t tornus (Fig. 4) (un like all other Ameletidae). Hin d wing has shape and size u sual for Am eletidae -- trian gular, w ith len gth slightly less th an 1/2 of fore w ing len gth . Su bim aginal win gs u nicolour. Egg. Un know n. Size. Small: length of mature larva 7 m m. DISTR IBUT ION . Known as a sin gle specim en from a single locality in R ussian Far East.


Review of Ameletidae of Russia and adjacent lands

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2 1

3

4

5
Figs 1-5 -- Metreletus spp. : 1 -- M. ba lcan icus, male larv al genita l b uds, v ent ral view. 2-5 -- M. micus ( holoty pe) : 2 -- ma le la rva l g enital buds, ve ntr al vie w; 3 -- la brum; 4 -- subim aginal fo re wing e xtr act ed fro m larv a; 5 -- te rga lius VI. Рис. 1-5 -- Metreletus spp. : 1 -- M. ba lcan icus, зач атк и генит алий л ичинки са мца , в ент рально. 2-5 -- M. micus ( голо тип): 2 -- з ача тки ге нит алий л ичинки самца, ве нтр ально; 3 -- в ерх няя гу ба; 4 -- с убимагина льное пере дне е к рыл о, изв леч енное из личинки; 5 -- т ергалия VI.

II. Genus Amele tus Eaton , 1885
H i e r a r ch i c a l n a me : A me l e t u s / g 2 ( s i n e M e t r e l e t us ; i n c l. Ch im u ra , Pa le o a m e l e tus ) REFEREN CE. Klu ge, 2 00 4: 82 , Fig. 20A ( lar va , ima go).

CHARACTERISTICS. Till recently, Ameletus was separated from Metreletus by the following characters: (1) Cubital field of fore wing retains several (4-8) veins going from CuA to basitornal margin. (2 ) Larval claws h ave no denticles. (3) Glossae truncate; all apical setae are flat, widened toward apex, rounded apically and form a single regular apical-ventral row. Now th e followin g character can be added: (4) Larval penis has simple sh ape: each lobe is triangular or conic, with rounded apex and without incision between lateral lobe and ventral plate (Figs 10 , 2 1); apex of this larval penis lobe correspon ds to the apex of im agin al lateral lobe. It is important to n ote, th at this sh ape of larval penis is retained n ot on ly in species whose imaginal lateral lobes are long, bu t also in A. oregon ensis, whose imaginal lateral lobes are shorter th an m edian plates and strongly divergen t. C OMP OSIT ION AN D D ISTR IBU TION . A meletus is distributed in H olarctic and Oriental R egion. It includes the followin g species:

O ne T ran s-H olarctic species -- A . ino pina tus E aton, 188 7 (= A. alp inus B engtsson, 191 3 = A. eug enii Sinitshen kova & Varykh anova, 198 9) (see below ). 33 other North American species (Zloty, 1996) -- A. amad or Mayo, 1939; A. andersoni Zloty, 1996; A. bellus Zloty, 1996; A. browni McDunnough, 1933; A. celer McDunnough, 1934 (= A. alticolus McDunnough, 1934 = A. celeroid es McDunnough, 1934 = A. tuberculatus McDunnough, 1939); A. cooki McDunnough, 1929; A. cryptostimulus Carle, 1978; A. dissitus Eaton, 1885; A. dodd sianus Zloty, 1996; A. ed mund si Zloty, 1996; A. exquisitus Eaton, 1885; A. falsus McDunnough, 1938; A. imbellis Day, 1952; A. lineatus Traver, 1932; A. lud ens Needham, 1905; A. majusculus Zloty, 1996; A. minimus Zloty & Harper, 1999; A. oregonensis McDunnough, 1933; A. pritchardi Zloty, 19 96; A. qu ad ratus Zloty & Harper, 199 9; A. shepherdi Traver, 1934 (= A. querulus McDunnough, 1938); A. similor McDunnough, 1928 (= A. monta Mayo, 1952); A. sparsatus McDunnough, 1931 (= A. aequivocus McDunnough, 1934); A. subnotatus Eaton, 1885; A. suffusus McDunnough, 1936; A. tarteri Burrows, 1987; A. tertius McDunnough, 1938; A. tolae Zloty, 1996; A. valid us McDunnough, 1932; A. vancouvernensis McDunnough, 1933 (= A. facilis Day, 1952);


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Nikita J. Kluge
Djako nov ) -- 1 Im #; Khibiny, lake Vud-Yavr , 1 930-19 37 (Frid olin) -- m any im ago es and sub ima goe s. KOM I REPU BLIC, No rthern and S ubp ola r U rals ( E. Nov iko va et al. ) -- ma ny LS- I, lar vae , subimag oes and imag oes. TUME NSKAYA OBLAS T' (= Tumen' P rov .): ba sin of river Vo ika r, 19. VIII.1 925 (Frid olin) -- 2 Im #; riv er Malaya So s'v a, 19 85 (T. Za guz ova ) -- 1 L; Ve rkhne- Taz ovskiy Na tur al Rese rve , r ive r P oko l'ka, 198 8 ( T. Za guz o va) -- 6 L . TAYMYRS KIY AUTON OMY ORRU G ( Ta ym yr Aut onom ous Pr ov .) : rive r Ko tuy, m outh o f river Ekhilakh, 2 6.VII. 197 9 ( N. Kluge) -- 1 larv al exuvia e; NW of Plato Putor ana, 22 .VII.1 982 (O. Antr opo va) -- 5 L. KH ABAROVSKIY KRAY (= Khaba rovsk P rov .), Tuguro-Chum ikanskiy Reg ion, r ive r U da, 4-5 km ab ove mo uth, 2 7-2 8.VIII.2000 (T. Tiunova ) -- 3 Im # ( dep osited in Vlad ivo sto k). MAGADANS KAYA OBLAST' (= Ma gad an Pro v.) , r ive r Yama , 2 1.VIII.19 80 (I. Zasy pkina) -- 9 Im #. MO NGO LIA: nort her n shore of la ke Hubsugul (K. Vary kha nov a) -- ma ny ima goe s a nd lar vae [see Sinit she nko va & Varyc hanova , 1 989 as A. eu genii].

A. velox Dodds, 1923 (= A. connectus McDunnough, 1936 = A. connectina McDunnough, 1939); A. vernalis McDunnough, 1924; A. walleyi Harper, 1970. Several species are kn own from C entral Asia -- A. primitivus Traver, 1 939; A. dod ecus (Dubey, 1 971 [Ichthybotus]) an d a group A. alexand rae (see below ). Th ree species are know n from Taiwan only -- A. atratus Kang & Y ang, 19 94; A. formosus Kang & Y ang, 1 994 and A. mo ntivagus Kang & Yang, 199 4. Four species are kn own from Japan on ly -- A. aetherea (N avas, 1 915 [Ch imura]); A. cro cerus Imanishi, 1 931; A. kyotoensis Im anishi, 19 31 and A. su balp inus Imanishi, 193 1. Several other species and subspecies are distribu ted in eastern part of Asia (see below) -- A. in opin atus la biatus Sin itsh en ko va, 1981 ; A . ca m tscha ti cus U lm er, 1927 (= A. pila tus Sinitshenkova, 198 1); A. altaicus sp.n.; A. mon tanus mo nta nus Im anishi, 19 30; A. mon tan us rossicus sub sp.n.; A. mo nta nus arlecchino sub sp.n.; A. pa rvus Kluge 197 9; A. ced ren sis Sinitshen kova, 1 977; A. costalis (Matsumura, 19 31 [Ch imura]) (= A. lon gulus Sinitshenkova, 1981 ). B elow , th ere are described th e species foun d on the territory of the former USSR (Russian Federation, republics of Central Asia an d C aucasus).

II.1. Ameletus inopinatus Eaton 1887
= A. a lpinus Bengtssonб 19 13 (synonymized with A. ino pina tus by Brekk e [ 19 65 ]) = A. lab ia tus Sinitshenk ova б 1 981 = A. eug e nii Sinitshenk ova & Var ycha nova б 19 89 (synonymized with A. in op in atus by Kluge [ 19 95 ]) REFERENCES. See separ ately for A. i. ino pin atus a nd A. i. la b i a t us .

La rva. Labru m length 0 .76- 0.8 2 of width (9 specimens m ea su re d) ( un like s h ort er in A . a lta icus an d long er in A. p arvus). Abdomin al terga and stern a w ithou t spine-like setae (un like group A. alexand rae). Tergalii I an d II w ithout an al ribs (un like A. costalis, A. formosus an d A. atratus); tergalius I smallest, tergaliu s II varies am ong su bspecies (see below). T ergalii III-VII w ith anal rib on an al margin (th e same in A. ca mtschaticus). Su bimago. Wings light brow nish, u nicolou r. Imago. Wings n on-coloured. M ale gen itals. In im ago an d su bim ago, lateral penis lobes ben t S-sh apely (u nlike all other species). In imago, ventral plates n on-developed, den ticles absent. Su bim agin al an d larval penis withou t spine-like setae. Egg. E llipsoid (not flattened). R eticulation: each polygonal an d roun d cell bears distinct papilla; all cells and papillae similar, so egg looks as even ly reticulated. Size. Medium: fore w ing len gth 9-1 1 m m. DISTRIBUT ION. Selected areas of H olarctic (see below, A. i. ino pin atus an d A. i. lab iatus). Su bspecies. T he species A. in opin atus can be divided in to 2 su bspecies (or form s) w hich differ only by len gth of larval 2nd tergaliu s. Areas of the both forms overlap in the Ru ssian Far East. Similar form s exist in A. mon tanus (see belo w ).

La rva. T ergaliu s II not elongated, n ot longer (either equ al or shorter) than tergalius III an d n ext ones. In most specimen s from Siberia and U rals, abdomin al darken ings and blan ks are con trastin g and distinctly outlined, bu t in some specimens they are non-contrasting and diffuse, as in specim ens from Great Britain [Macan, 1961 : Fig. 17d]. Su ch larvae with non-contrasting coloration can be confused with A. ca mtschaticus. Subimago, imago, male genitals and egg. See A. inopinatus. D ISTR IBU TIO N. Selected areas of Holarctic: in Europe -- Great Britain, Ju ra Moun tain s, Alps, Carpathians, Scan din avia, Kolskiy P en insu la, north of Ural Moun tains; in Asia -- Mou ntain s Pu toran a (to th e south from Taimyr P en insu la), n orthern tributaries of Lake H u bsu gu l (= H цvsgцl Nu ur) (in Mongolia), mountains to the north-vest from Sea of Okhotsk; in N orth Am erica --Alaska and Nothwest Territories of C anada [Zloty, 1996 ].

II.1.b. Ameletus inopinatus labiatus Sinitshenkova, 1981
= Am e letu s lab ia tus Sinitshenk ova , 1981 = Am ele tus p roc erus: Klu ge, 198 2 ( non Am ele tus pro cer us Ba jk ova , 1 976) RE FE REN CE S. Sin it she nk ova , 19 8 1 : 7 5 , Fig . 2 ( la r va ); Kl ug e, 1 98 2 : 1 1 3 , Fi g. 2 ( ma l e ima g o, su bi ma go , l a r va ; as A. proc erus); Tsher nova et a l., 19 86: 12 6, Figs 5 7: 9, 10 (ma le imago, as A. pr oc erus); Kluge, 1 995 : 9 , Figs 2 -3 (la rva ). MATERIAL EXAMIN ED. RU SSIA: BURYATIYA: 13 .VI.2002 (V. Ivanov, S. Melnit sky ) -- 1 L/S #, 1 L $; riv er S nez hna ya (sout her n t rib uta ry of Lake Baikal) , 2 4.VI.1 998 (N . Rozhkov a) -- 2 L ; natural reserv e Barg uzinskiy: river Dav sha , 1 km fr om La ke Baikal, 2 9.VI.19 98 (N. Ro zhkova , O. Rusinek) -- 1 L ; ibid ., mo uth of river Osinov ka, 22 .VII.1 998 (N . Ro zhkova ) -- 2 L; na tur al reserv e Ba ika lskiy, river Ma linovka, 2 4.VII. 199 8 ( N. Ro zhkova ) -- 1 L. KHABAROVSKIY KRAY (= Khaba rov sk Pr ov. ): riv er Nikita , 1 0 km N railwa y stat ion Bira, 4. VI. 198 0 ( N. Kluge ) -- 2 L-S -Im #; 50 L. PRIMORSKIY KRAY, na tur al re ser ve Sikhote-Alinskiy : stre am nea r Terney, 14. IV. 197 4 ( L. Na dez hdina) -- 1 L ( holoty pus of A. la biatus); river Yasnay a, 19 .IV.19 74 (L. Na dez hdina) -- 1 L ; r ive r Zabo loc hennay a, 18 .IV.19 74 (L. Na de zhd ina ) -- 1 L.

II.1.a. Ameletus inopinatus inopinatus Eaton, 1887
= A. alpin us Bengtsson, 1 913 = A. e u ge n ii Sinitshenk ova & Va r ycha nova , 1989 REFEREN CES. Sinitshenkova & Va r ycha nova , 198 9: 5 76, F i g s 1- 2 0 ( m a l e a nd f e ma l e i m a go , s u b i ma g o , l a r v a ; as A. eu genii); Zloty & Pritchard, 1 997 : 2 61, Figs 1, 6, 18A, 2 2E ( l a r va , i ma go ). M AT E R IA L E X A M INE D . R U SS I A: M U R M A N S KAYA OBLAS T' (= Murm ansk P rov .): Ko lskiy Zaliv, 28. VI. 191 5 ( A.

La rva. T ergaliu s II elongated, longer than tergalii III-V II [Kluge, 1 982: Fig. 2: 4 ]. Abdom inal terga and stern a usually with contrasting darken ings an d blanks, so that dark oblique stripes are indistin ct or non -expressed on backgroun d of darkenings. Subimago, imago, male genitals and egg. See A. inopinatus. DISTRIBUTION. From Transbaikalia to Russian Far East. CO MME NT. Klu ge [19 82] described this species under th e n ame A. pro cerus B ajkova, 197 6, basing on a wrong mu seu m label "h olotype" w hich was placed by B ajkova on


Review of Ameletidae of Russia and adjacent lands
th e specimen which is actu ally paratype. T he true holotype of A. pro cerus belongs either to A. ced rensis, or to A. montanu s rossicus; the name A. pro cerus should be regarded as nomen dubium (see below ).

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II.2. Ameletus camtschaticus Ulmer, 1927
= Ame le tu s p ilatus Sinitshenkova , 1 98 1 (synonymized with A. ca mtsc ha ticus by Kluge [1 99 5]) REFERENCES. Ulmer , 1 927 : 12 , Figs 10 -12 (male imago); Sinitshenkova & Tshernova, 1 976 : 13 , Figs 3- 9 (male su bimago, larva); Sinitshenkova, 198 1: 73, Fig. 1 ( lar va, as A. pilatus); Tshernova et a l., 1 986 : 126 , Figs 57:7- 8 ( ma le imago); Kluge, 2 00 4: Figs 1 1:A-D ( genitals of ma le imago). MATERIAL EXAMIN ED. RU SSIA: KHABAROVS KIY KRAY (= Khaba rov sk Pro v.) , 6 km E Obluch'ye , 2 8.VII-3.VIII.19 84 (N . Klug e) -- 3 L-S -Im #, 1 L -S /Im #, 4 L -S -Im $, 1 L -S $. PRIMORSKIY KRAY, S ikhote -Alin' Na tur al Rese rve : stre am Ke d r o v y i, 3 0. VI- 6 . VII. 1 982 ( E . P o t ikha , L . N e s t e r o v a , I. So han') -- 7 L/S, 1 L; st rea m Yasnyi, 12 .VI.198 1 ( E. Pot ikha) -- 3 L /S; st rea m S por nyi, 1 3.VI.19 90 (E. Po tikha) -- 2 S -Im $, 5 S $, 3 S #, 1 L /S $, exuvia e L L. MAGADANSKAYA OBLAS T' (= Ma gad an Pro vinc e): river Khasy n near Khasyn, 2 8.VIII.19 78 (V. Zher ikhin, N. Sinitshenkov a) -- 1 Im #; nea r S ibit- Tellakh, st rea ms Ole n' and Osernyi, 2-2 1.VII. 197 7 ( L. Zhiltz ova ) -- 2 Im #, 1 Im $, 7 S #, 20 S $, 4 L /S, 8 L.

Larva. Labrum length 0.78-0.81 of width (5 specimens measured) (unlike shorter in A. altaicus and longer in A. parvus). Abdominal terga usually withou t contrasting darkenings, with a pair of diffuse dark oblique su bmedian stripes an d diffu se blanks. Abdominal terga an d stern a w ithout spin e-like setae (u nlike group A. alexand rae). Tergalii I and II withou t an al ribs (unlike A. costalis, A. formosus an d A. atratus), sm aller th an others. Tergalii III- VII with an al rib on anal margin (the sam e in A. in opin atus). Su bimago. Wings brown ish , u nicolou r. Imago. Wings n on-coloured. Ma le g enitals. In im ago, lateral penis lobes with apices moderately bent medially an d n ot bent dorsally; each ventral plate bears a large apical den ticle and several sm all den ticles proximad of it. Subimagin al penis w ith the same number of spine-like setae: one pair of large sclerotized ones an d several sm aller ones proxim ad of each large one. Larval penis with on e pair of spin e-like setae (which correspond to large on es in subimago), rarely w ith one more pair of smaller spine-like setae proxim ad of them. Egg. E llipsoid (not flattened). R eticulation: each polygonal cell with flat bottom lacking protuberan ce, with borders lower and narrow er than borders of roun d cells, so round cells (each bearin g papilla) are promin ent; round cells small and sparse on most surface, denser and larger on pole. Size. Medium: fore w ing len gth 9-1 2 m m. DISTRIBUT ION . Ru ssian Far East.

II. 3. Amele tus alt aicus Kluge, sp.n. (Figs 6-12, 17)
REFEREN CES. See sepa r ately for A. m . m o ntan us a nd A. m . r o ss ic us s ubs p.n. MATERIAL EXAMINED. RUSSIA: ALTAY: 10 km N KoshAgach, mountain stream running from ridge Kurayskiy, 4.VIII.1987 (N. Kluge) -- L-S-Im #(holotypus); ibid., 2-5.VIII.1987 (N. Kluge) -- 3 L-S-Im #, 3 L-S-Im $, 4 L; river Tekelushka near its falling to river Kucherda (tributary of Katun'), 24-25.VII.1996 (S. Melnitsky) -- 1 L. TUVA, river Balyktyg-Uem, 26.VI.1996 (V. Zaika) -- 1 Im #; IRKUTSKAYA OBLAST' (= Irkutsk Prov.): Bolshie Koty, rivers Sennaya and Kotinka (vestern tributaries of Lake Baikal), 9-11.VIII.1994 (N. Kluge) -- 9 L-S-Im #, 1 L-SIm $. BURYATIYA: natural reserve Baikalskiy, river Kurkavka, 23.VII.1998 (N. Rozhkova) -- 1 L; river Snezhnaya (southern tributary of Lake Baikal), 24.VI.1998 (N. Rozhkova) -- 2 L.

La rva. C uticle light, with non-contrasting, diffu se blan ks an d darkenin gs. Head capsu le and clypeus with diffuse darken ings. Labrum shorter than in other species, its len gth 0.5 9- 0.68 of w idth (6 specimen s measu red). Labrum with diffu se darken ing in a form of triangle or a pair of oblique ben ds w h ich stretch from an terior-m edial in cision to posteriorlateral corn ers. Mandibles can h ave more or less expressed diffu se distal and proximal darken ings. Thorax light, with diffuse ornament of blanks an d darkenin gs. Legs m ostly ligh t, with darkened apex of tarsus and som etimes with m ore or less expressed darkenin gs on join ts. Abdominal terga ligh t, with darkenin gs non-con trastin g and diffuse; most terga have un iform pattern of a pair of diffuse dark obliq ue submedian stripes and diffuse blanks (Figs 6-7 ). Abdominal n erve ganglia as in imago. Abdom inal terga an d stern a without spin elike setae (u nlike group A. a lexa nd rae). T ergalii I an d II w ithou t an al ribs (un like A . co s ta lis, A . fo r m o sus and A . a tratus), sm all. T ergalii III- VI with an al rib separated from anal margin (un like A. in opin atus an d A. ca mtschaticus), bu t located near an al margin (un like A. mon tanus and oth er species); tergalius VII with anal rib ju st on anal margin (Fig. 1). Caudalii either nearly unicolour, or with diffu se colour pattern -- proxim al blan k, m iddle darkening, distal blank and darken ing on extreme apex. Su bimago. Wings brown ish , u nicolou r. Imago, male. H ead brown. Thorax: sclerites brown (from dark to ligh t brown ), membran es pale; in each specim en all sclerites (n otum , pleu ral sclerites and sternu m) have similar colour (un like A. mon tanus). Fore legs brow nish. Middle and hind legs pale; coxa, troch anter and articulations brownish. Wings with m embrane colourless, vein s brow nish ; pterostigma withou t distinct coloration. Abdomen: Segm ent I brow n. Segm en t II in most part light brown ish , posterior margin pale. Segmen ts III- VI tran slu cen t, in m ost part pale; on each of them tergum w ith diffuse brow nish stripes alon g lateral m argin s, diffu se brow n ish su bm edian stripes an d diffu se brownish transverse ban d parallel to posterior margin (posterior margin pale). Segmen ts VII-VIII n on-tran slucent, with th e same brow nish m aculation. Segments IX-X darker. Last abdominal nerve ganglion dark brown, oth er ganglia either as pale as sterna, or partly brow n. Cerci pale, sometimes with brow nish articu lation s. Ma le g enitals. In im ago, lateral penis lobes with apices strongly ben t medially and somew hat dorsally; each ventral plate with several (4-1 0) equ ally sm all den ticles (Figs 9, 1 1, 12 ). Subimaginal penis with the sam e n umber of spine-like setae w h ich correspon d to th e im agin al den ticles. Larval penis with less num ber (0- 5) of spin e-like setae (Fig. 10 ). Imago, female. H ead pale. Thorax with sclerites pale or light brownish. All legs pale. Wings with membran e colou rless, veins brow n ish ; pterostigma w ith out distinct coloration, with anastomosed veins. Abdomen either en tirely pale, or with brow nish pattern on terga an d ganglia pigmen tation as in male. Cerci pale, sometimes w ith brownish articulation s. E gg . E llipsoid (n ot flatten ed). R eticu lation : polygonal cells with bottoms eith er flat or bearin g a small protuberance, with borders promin ent; round cells eith er absent, or a few large prominent rou nd cells (each bearin g papilla) exist on one pole only (Fig. 17 ). Size. Medium: fore w ing len gth 9-1 0 m m. D ISTR IBUT ION . Siberia, at least from Altay Mountains to Baikal Lake. CO MPAR ISO N. T he new species resem bles North American species A. celer in structure of m ale imagin al genitals, larval cu ticular colou r pattern of abdomen and position of


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Nikita J. Kluge

6

7

8

10

12

9

11

Figs 6-1 2 -- Am eletus a lta icus, sp.n., holoty pe: 6 -- larv al ter ga I-II with right ter galii I-II; 7 -- la rva l t erg a VI-VIII with right te rga lii V-VII; 8 -- male larv al genita l b uds, v ent ral view; 9 -- ma le ima ginal genita ls, ve ntr al vie w, the sa me mag nifica tio n; 10 -- enlarg ed male larv al penis bud s, ventra l v iew; 1 1 -- ma le imag ina l p enis, ventra l v iew, t he sam e m agnificat ion; 1 2 -- ma le im aginal pe nis, la ter al vie w. Рис. 6-1 2 -- Am eletus a lta icus, sp.n., гол отип: 6 -- личиночные тер гит ы I-II с прав ыми те рга лия ми I-II; 7 -- личиночные те ргиты VI-VIII с пр авыми тер гал иями V-VII; 8 -- за чат ки генита лий личинки самца, ве нтр ально; 9 -- генитал ии има го са мца , в ент рал ьно , т о ж е у вел иче ние ; 1 0 -- ув еличенные за чатк и пениса лич инк и с амца, вентра льно; 11 -- пенис имаго с амца, ве нтр ально, то же ув еличение; 12 -- пенис имаго с амца, лат ера льно.


Review of Ameletidae of Russia and adjacent lands
an al rib on tergalii [Zloty & Pritch ard, 19 97: Figs 4 , 17 B, 22 C]; unlike A. celer, the new species h as eggs not flattened.

253

II.4. Ameletus montanus Imanishi, 1930 (Figs 13-16, 18-19)
Larva. Labrum length 0.78-0.92 of width (9 specimens measured) (unlike shorter in A. altaicus and longer in A. parvus). Abdominal terga and sterna usually w ith contrasting darkenings and blanks (varying among subspecies -- see below), so that dark oblique stripes indistinct or non-expressed on background of darkenings. Abdom inal terga and sterna without spine-like setae (unlike group A. alexand rae). Tergalii I and II without anal ribs (unlike A. costalis, A. formosus and A. atratus); tergalius I smallest, tergalius II varies among subspecies (see below). Tergalii III-VI with anal rib well separated from anal margin (unlike A. inopinatus, A. camtschaticus and A. altaicus); only tergalius VII with anal rib on anal margin. Su bimago. Wing colour varies am ong subspecies (see belo w ). Ma le g enitals. In im ago, lateral penis lobes with apices straight in ven tral view, ben t dorsally; each ven tral plate with several (1-7) eq ually small denticles (Figs 15- 16). Su bimagin al penis w ith the same n umber of spin e-like setae corresponding to the imagin al denticles. Larval pen is with less nu mber (0 -3) of spine-like setae (as in Fig. 10 ). Egg. Flattened, in a form of ellipse-like biconvex lens (Figs 18-19). Reticulation: each polygonal cell with flat bottom lacking protuberance, with borders much lower and narrower than borders of round cells, so round cells (each bearing papilla) are prominent; round cells small and sparse on most surface, dense and large on one pole (the same in A. cedrensis and A. costalis). Size. V ariable: from 7 to 1 0.5 mm (see below ). DISTRIBUT ION . Altay, East Siberia, R ussian Far East, J apan. CO MME NT. Am eletus pro cerus B ajkova, 19 76 was synon ymized with A. mon tanus by Kluge [1 995]. Actu ally, the holotype of A. pro cerus can belong either to A. ced rensis, or to A. mon tan us rossicus, so the species n ame A. procerus sh ould be regarded as nom en dubium (see below ). Su bspecies. T he species A. mon tanus can be divided in to th ree subspecies (or forms) wh ich differ mainly by len gth of larval 2n d tergalius (the same in two su bspecies of A. inop inatus -- see below ) and coloration of subimaginal w ings (the same in form s of group A. alexand rae).

M A TE R IAL E XA M IN E D . RU SS I A: A L TA Y, Tu ra c ha k, r ive r Biya , 1 0-3 1 . VII. 1 937 ( L . Go r b uno v ) -- 1 L /S # , 1 L. BU RYATIYA: nat ura l r e se r ve Ba ika lskiy , r ive r P e r ee m naya, 23 . VII. 1 998 ( N . Ro z hko v a) -- 11 L /S . P RIM ORS KIY KRAY: nat ura l r e se r ve Ke d r o v aya Pa d ', str ea m Bo lsho y Zo lo t o y, 22 -23 . VII. 1 98 0 ( N . Kluge ) -- 1 L - S - Im # ( ho lo ty pus ) ; ib id, 2 0-26 . VII. 1 98 0 ( N . Kluge ) -- 1 L - S - Im # , 1 L - S - Im $, 5 L/ S ; r ive r N ar v a ( = S id im e ) , 2 0-2 1 . VII. 1 98 0 -- 2 L - S - Im # , 1 L -S #.

II.4.a. Ameletus montanus montanus Imanishi, 1930
REFERENCE. Ima nishi 19 30 : 2 65 , Figs 3 -5. MATERIAL EXAMIN ED. RU SSIA: KURIL ISLANDS , S hikota n, Malo-Kurilsk, 1 4.VII.1 971 (E rmo lenko) -- 1 S #. JAPAN: Na gano P ref., Kom aga ne Cit y, N aka gosha Valley , H igy rashi wa ter falls, 24 .IX.2002 ( I. Tat are nko ) -- 1 L/S #, 2 L /S $, 4 L.

Larva. Tergalii II not elongate, oval, smaller than tergalii III (as in A. m. rossicus subsp.n., unlike A. m. arlecchino subsp.n.). Su bimago. Wings brown, crossvein s bordered by darker brown. Egg. U nknown; probably, as in other A. mon tanus (see above). Size. Mediu m: fore wing length 9 -10 .5 m m. DISTRIBUT ION . Japan an d Ku ril Islands.

II.4. b. Amele tus mont anus rossi cus Kluge, subsp.n.
= A m e le tu s m o n t a n us : B a j ko va , 1 9 7 6 ; Si ni ts he nk ov a & T she r nova , 1 976 ; T sh er nova et a l., 1986 REFEREN CES. Ba jk ova , 1976 : 582 , Figs 1-7 ( ma le a nd fe ma le ima go a nd la r va ) ; Si nit shenk ov a & T sh er no va , 19 7 6: 17 , Figs 18 -21 (male su bimago, la rva ); Tshernova et al., 198 6: 12 6, Figs 5 7: 5, 6 (male imago).

La rva. T ergalii II not elongate, oval, sm aller than tergalii III (as in A. m. mon tanus, unlike A. m. arlecchino sub sp.n.). C u ticle often h as con trastin g blan ks an d darken in gs, bu t u s u ally n ot s o p ronoun ce d a s in A . m . a rl ecch i no su bsp.n. H ead capsu le m ostly darken ed (exce pt blan ks on eyes an d ocelli) ; clypeu s c om p let ely dar kene d. Labr um coloration var iable -- eith er m ostly ligh t w ith n on -contrastin g darken in g basally, or w ith m ore or less con trasting distal-m edian b lan k an d proxim al-lateral da rken in g (B ajkova, 1976 : Fig s 4-6 ; Sin itsh en kova & T sh ern ova, 1976: Fig . 1 9 ), o r m os tly da rk en e d. M an dible w it hou t d is tal darken in g: w h ole distal h alf is occu pied by blan k, darkenin g presen t on proxim al part on ly (like A . m . a r lecch ino sub sp.n., un like othe r species). Maxilla w ith darken in g on proxim al part o f stipes. An tenn al pedicellu s partly darken ed, flagellu m darker in m iddle part, ligh ter proxim ally an d apically. T h orax w ith orn am en t of blan ks an d darkenin gs. Legs m ost ly ligh t, w ith darken ed apex of tarsu s and w ith m ore or less expressed oth er darken in gs -- on coxa, on troch an ter ( or on troch an tero-fem oral jo in t), on m iddle of fem u r, on ba se of tibia, on base of tarsu s (or on tibiotarsal join t). Abdom in al cu ticle h as colou r pattern u su ally w ith 4 m iddle t erga III- V I distin ctly darke r th an previous an d n ext one s [B ajkova, 1976 : Fig. 7 ] (un like A . m . a rlec ch in o su bsp.n., lik e A . ce d ren sis an d som e oth ers ): T ergu m I m ostly darken ed. T ergu m II m ost ly occu pied by blan k, w ith a sm all m edio-an terior darken in g. T ergu m III eith er also m o stly ligh t, or w ith som e ot h er darken in gs resem blin g ter ga IV - V I. T erga IV - V I m ost ly occu pied by darken in g, w ith a pair of isolated, large o r sm all blan ks, som etim es w ith isola ted, un paired blan k betw een th em. T er g a V II- V III m os t ly oc cu p ied b y b lan k , w it h s m all darken in gs. T e rga IX - X m ostly occu pied b y darken in gs, w ith sm all blan ks. Stern a I- II m ostly or co m pletely occupied by blan k. Stern a III- V I eith er th e sam e, or w ith some darken in gs. St ern a V II- V III, w ith m ore or less expressed m edian darken in gs an d lateral blan ks. Ste rnu m IX m ostly occu pied by da rken in g. Abdom in al n erve g an glia n on -colou red. C au dalii w ith proxim al blan k (of va riable len gth ), m iddle darken in g (term in atin g distad of m id len gth ), distal blan k an d darken in g on extrem e apex. For other ch aracters -- see A. mon tanus. Su bimago. Wings un iformly brown (un like A. m. montanus). Imago, male. H ead brown. Thorax: Pleural sclerites and sternu m brown , pleu ral mem bran es pale; m esonotu m lighter th an pleu rites and sternite, only an telateroparapsidal sutu re brown. Fore legs brownish. Middle an d hind legs pale; coxa, trochanter and articulations brownish. Win gs w ith membrane colourless, vein s brow nish ; pterostigma tinged with brow nish, w ith crossveins in distal parts of costal and subcostal fields n arrowly bordered by brow n. Abdomen: Segment I brown . Segments II-VI translucent, in most part pale; on each of them tergum w ith diffuse brownish stripes along lateral m argins con nected with diffu se brownish tran sverse ban d parallel to posterior m argin (posterior m argin pale); sterna either un iformly pale, or with diffu se m edian brow nish makin gs. Segmen ts V II- IX n on-translu cen t, w ith th e same


254

Nikita J. Kluge II.4. c. Amele tus mont anus arlecc hino Kluge, subsp.n. (Figs 13-16, 18-19)
MATERIAL EXAMINED. RUSSIA: Primorksiy Kray, Terney and Sikhote-Alinskiy natural reserve: Terney, river Serebryanka, 18.VIII.1990 (N. Kluge) -- 1 L-S #, 1 L/S #; river Maysa (= Yasnaya), 4.VIII.1990 (N. Kluge) -- 1-L-S-Im #(holotypus); ibid., 9-15.VIII.1990 (N. Kluge) -- 8 L-S-Im #, 3 L-S-Im $, 30 L; ibid., 14.VII.1982 (E. Potikha) -- 1 L/S #, 1 L; river Zabolochennaya, 13.VII.1982 (E. Potikha) -- 1 L/S #; stream Kedrovyi, 13.VIII.1990 (N. Kluge) -- L-Im #; Blagodatnoe, VII.1990 (N. Kluge) -- 1 L/S #.

brown ish maculation . Segm ent X darker. Abdominal nerve ganglia n on-coloured. Ma le g enitals. See A. mon tanus. Imago, female. H ead pale. T horax as in m ale. All legs pale. Win gs as in m ale. Abdomen pale, with diffuse markin gs as in male. Egg. See A. mon tanus. Size. Mediu m: fore win g length 8 -9 m m. DISTR IBU TION. Altay, E ast Siberia an d R ussian Far E ast.

13

14

15

16
Figs 13-1 6 -- Am eletus mo ntan us arlecc hino, sp.n.: 13 -- larv al ter ga I VIII wit h r ight t erg alii V-VII; 15 -- m ale im aginal ap ex of abd ome n with ve ntr al vie w ( 13-15 -- holoty pe) . Рис. 13-1 6 -- Am eletus mo ntan us arlecc hino, sp.n.: 13 -- л ичиноч ные личиночные тер гит ы VI-VIII с пра выми т ергалиями V-VII; 15 -- вер шин 16 -- генита лии има го самца, ве нтр ально (13-15 -- голо тип). -II with rig ht ter galii I-II; 14 -- la rva l t erg a VI- ge nit als, late ral view; 16 -- male imag ina l g enitals, те ргиты I-II с пра выми т ергалиями I-II; 14 -- а брюш ка има го самца с генитал иями, лат ера льно;


Review of Ameletidae of Russia and adjacent lands
La rva. T ergaliu s II elongated, longer than tergalii III-V II (un like A. m. mon tanus an d A. m. rossicus sub sp.n.). Cu ticle often h as contrasting blanks and darkenings. Head capsule m ostly darkened (except blan ks on eyes and ocelli); clypeus completely darkened. Labru m h as distal 1/3 completely occu pied by blank; proximal 2 /3 either com pletely occupied by contrasting darken ing, or with pair of con trasting darkenings separated by a m edian longitudin al blank. Mandible without distal darkenin g: w hole distal h alf is occupied by blan k, contrasting darken ing is lim ited by proxim al h alf (like A . m. ro ssicu s sub sp .n., un like oth er species). Maxilla w ith darken ing on proximal part of stipes. An tenn al pedicellu s partly darkened, flagellum darker in middle part, lighter proximally and apically. Thorax with composite orn am en t of contrastin g blanks an d darken in gs; in last larval in star, each fore protopteron has darker proximal part (which overlaps dark abdomin al tergu m I) and ligh ter apical part (w hich overlaps light abdomin al terga II-III). Each leg mostly occupied by blan k, with several contrastin g darkenin gs (which can be larger or sm aller) -- on coxa, on troch anter (or on trochantero-femoral joint), on m iddle of femur, on base of tibia, on base of tarsu s (or on tibio-tarsal joint) an d on apex of tarsus; claw non -darken ed. Abdominal cuticle has characteristic colour pattern, strongly different on different segments, usually w ith 3 m iddle terga IV -VI mu ch darker than previous and n ext on es (Figs 1 3-1 4): T ergum I m ostly darken ed. Tergum II mostly occupied by blan k, with a small medioan terior darkening. Tergum III either th e same, or with some other darkenings. T erga IV -VI mostly occupied by darkenin g, with a pair of isolated, contrasting, large (Fig. 14) or small blan ks, sometim es w ith isolated, u npaired blan k betw een th em. Terga VII-VIII m ostly occu pied by blank, w ith small darken in gs. T erga IX- X mostly occu pied by darken ings, with small blanks. Sterna I-II mostly or com pletely occupied by blank. Sterna III- VI eith er the same, or with m edian blanks an d lateral darkenings; from segmen t III to segment VI blanks become sm aller and darkenings larger. Stern a VII- VIII, vice versa, w ith median darkenings (sometimes nondeveloped) an d large lateral blan ks. Sternum IX mostly occupied by darkenin g. Abdomin al n erve ganglia non-coloured. Caudalii with contrasting proximal blan k (of variable length ), middle darken in g (term in ating distad of midlength), distal blank and darken ing on extreme apex. For other ch aracters -- see A. mon tanus. Su bimago. Wings un iformly brown (un like A. m. montanus). Imago, male. H ead brown. Thorax: Pleural sclerites and sternu m brown , pleu ral mem bran es pale; m esonotu m lighter th an pleu rites and sternite, only an telateroparapsidal sutu re brown. Fore legs brown ish or reddish . Middle and hin d legs pale; som etimes fem ur indistin ctly tinged w ith reddish near middle. Wings with mem bran e colou rless, veins brownish; pterostigma tinged with brownish, with crossveins in distal parts of costal an d su bcostal fields narrow ly bordered by brown. Abdomen: Segment I brow n. Segm ents II-VI translucent, in most part pale; on each of them tergum with diffu se brownish or reddish pattern -- pair of stripes alon g lateral margin s connected with tran sverse ban d parallel to posterior margin (posterior m argin pale); sterna either uniformly pale, or with diffu se median brownish m akin gs. Segmen ts VII- IX non-translucent, with the same brownish maculation . Segment X darker. Abdominal n erve ganglia n on-coloured. Ma le g enitals. See A. mon tanus. Imago, female. H ead pale. T horax as in m ale. All legs pale. Win gs as in m ale. Abdom en pale or with diffuse markin gs as in male.

255

Egg. See A. mon tanus. Size. Small: fore win g length 7 -8 m m. DISTRIBUT ION . Kn own from one area on th e eastern slope of Sikh ote-Alin' Moun tain s.

II.5. Ameletus parvus Kluge, 1979 (Figs 20-21)
REFERENCE. Klu ge, 1 97 9: 80 7, Figs 1- 15 (ma le a nd female imago, subimago, lar va). MATERIAL EXAMIN ED. KAZAK HS TAN: VOSTOCHNOKAZAKHSTANS KAYA OBLAST' (= East-Kaza khstan Pr ov. ), 90 km S Ust'-Kamenogo rsk, m ountain r idg e Kalbinskiy Khr ebe t, 6-8.VIII.19 78 (N. Kluge ) -- 2 L-S -Im #, 2 L -S -Im $, 26 L (holo typ us & pa rat ypi).

Larva. Labrum longer than in most other species, its length 1.0-1.03 of width (3 specimens measured). Abdomen nearly unicolourly darkened, without blanks and with indistinct dark oblique stripes. Abdominal terga and sterna without spine-like setae (unlike group A. alexand rae). Tergalii I and II without anal ribs (unlike A. costalis, A. formosus and A. atratus), small. Tergalii III-VI with anal rib well separated from anal margin (unlike A. inopinatus, A. camtschaticus and A. altaicus sp.n.); tergalius VII with anal rib nearer to anal margin. Su bimago. Wings brown ish , u nicolou r. Imago. Wings n on-coloured. Ma le g enitals. In im ago, lateral penis lobes with apices nearly straight in ven tral view, slightly ben t dorsally; ventral plates non-sclerotized, in distal part with a few (3 -4) small denticles (Fig. 20) (un like A. mon tanus, A. altaicu s sp.n. and A. camtscha ticus w hich have denticles in proximal part of ventral plate); these denticles are pale and poorly visible, so in the original description they were overlooked. Su bimagin al penis with the same num ber of den ticles or spin e-like setae correspon din g to im agin al ones. Larval penis with less nu mber (0 -1) of spine-like setae in middle part (Fig. 21 ) (unlike A. alta icu s sp.n., A. mo nta nus and A. ced rensis, whose larval penis has spine-like setae in proxim al part -- Fig. 10 ). Egg. Flattened, in a form of ellipse-like bicon vex lens (as in A. mon tanus an d A. ced rensis). Reticu lation: each polygonal cell with bottom bearing a protuberance (un like A. montanus an d A. ced rensis), with borders low er than borders of round cells, so rou nd cells (each bearing papilla) are prom inent; rou nd cells are small and sparse on most su rface, den se an d large on one pole. Size. Small: fore wing len gth 7 m m. DISTR IBUT ION . Altay (know n as type series from the single locality).

II.6. Ameletus cedrensis Sinitshenkova, 1977
REFER EN CES. S initshenk ova , 19 7 7 : 44 , F igs 1-2 ( ma le and fema le ima go, lar va) ; T shernova et al., 1 986: 12 6, Figs 5 7: 3, 4 ( ma le ima go). MATERIAL EXAMIN ED. RU SSIA: ALTAY: riv er Biya , 1 .VII- 25 .VIII. 193 7 ( L. Gor bunov) -- 40 L; riv er Koldor ne ar its fa lling into lake Telet sko e, 15. VIII.1 987 (N . Kluge ) -- 3 L-S -Im #, 4 L- S-Im $, 1 L -S #; nat ura l r ese rve Altay skiy: river Ko rbu ne ar it s falling into lake Tele tskoe, 19 -22 .VIII. 198 7 ( N. Kluge) -- 5 L-S -Im #, 2 L -S -Im $, 2 L -S #, 1 L -S $; ibid., Ya ily u, 13 .VIII. 198 7 ( N. Kluge ) -- 1 L-S -Im $, 1 L . IRKUTSKAYA OBLAS T' (= Irkutsk P rov .): Bolshie Koty , r ive rs Sennay a a nd Ko tinka (we ste rn t rib uta rie s o f L ake Ba ika l), 7-1 3.VIII.19 94 (N . Klug e) -- 20 L-S -Im #, 11 L-S -Im $, 3 L -S $. KH ABAROVSKIY KRAY (= Khaba rov sk Pr ov. ): 6 km E ra ilway st ation Obluch'e, railwa y stat ion "U dar nyi", 27.VII-3.VIII.19 84 (N . Klug e) -- 6 L-S -Im #, 3 L -S -Im $, 2 L -S #, 6 L. PRIMORSKIY KRAY: U pper-U ssuri resear ch stat ion of Biologic al- Soil Inst itut e, 35 km SE Chugue vka , 1 .VIII.1 980 (N . Klug e) -- 2 L -S -Im #, 12 L; ibid., 7 VII 19 75 (L . Zhiltzo va) -- 1 S #;


256
ib id. , 2 6-2 7.VI.19 75 -- 3 pa rat ypi); nat ura l r ese arc h 10.VIII. 198 0 ( N. Kluge ) -- 3 L-S $; ibid., river Yasnay S- Im #; str eam Sikho te, 21 na tur al resear ch Ked ro vay a

Nikita J. Kluge
Im #, 6 Im $, 6 L (holoty pus & S ikho te- Alinskiy, st rea m Kedr ovy i, 2 L-S -Im #, 2 L -S -Im $, 1 L -S #, a, 13. VIII.1 990 (N . Kluge ) -- 1 L.VIII. 19 79 (Timoshkin) -- 1 Im #; P ad', 1 Im #, 12 L ( pa rat ypi).

Larva. Labrum length 0.85-0.90 of width (3 specimens measured) (unlike shorter in A. altaicus and longer in A. parvus). Abdominal terga and sterna usually w ith contrasting darkenings and blanks, so that dark oblique stripes indistinct or non-expressed on background of darkenings. Abdominal terga and sterna without spine-like setae (unlike group A. alexand rae). Tergalii I and II without anal ribs (unlike A. costalis), small. Tergalii III-VI with anal rib well separated from anal margin (unlike A. inopinatus, A. camtschaticus and A. altaicus sp.n.); only tergalius VII with anal rib on anal margin. Su bimago. Wings brown ish , u nicolou r. Imago. Wings n on-coloured.

Ma le g enitals. In im ago, lateral penis lobes with apices straight in ven tral view , ben t dorsally; each ven tral plate projects far ven trally, w ith a sin gle lon g apical den ticle, withou t small denticles. Su bim aginal pen is also has on e pair of large den ticles correspon din g to im agin al on es. Larval penis has on e pair of spin e-like setae corresponding to imagin al and subimaginal denticles. Egg. Flattened, in a form of ellipse-like bicon vex len s. Reticu lation: each polygonal cell with flat bottom lacking protuberance, with borders mu ch lower and n arrower th an borders of round cells, so round cells (each bearing papilla) are prom in en t; rou nd cells are small and sparse on m ost su rface, den se and large on one pole (the same in A. mon tan us and A. costalis). Size. Mediu m: fore wing length 12- 14 m m. DISTR IBUT ION . Siberia (from Altay at the west) and Far East.

17

20

18

21

19
Figs 17-21 -- Ameletus spp.: 17 -- A. altaicus, sp.n., A. parvus: 20 -- male imaginal penis, ventral view; 21 Рис. 17-2 1 -- Am eletus spp. : 1 7 -- A. alta icus, sp.n., 20-21 -- A. pa rvus: 20 -- пенис имаго с амца, в ент рал egg; 18-19 -- A. montanus arlecchino, subsp.n., egg in two views; 20-21 -- -- male larval penis buds, ventral view, the same magnification. яйцо; 1 8-1 9 -- A. mo nta nus arlecc hino, subsp.n., яйцо в ра зных р аку рса х; ьно ; 2 1 -- за чат ки пениса личинки самца, ве нтра льно, то же уве лич ение.


Review of Ameletidae of Russia and adjacent lands II.7. Ameletus costalis (Matsumura, 1931 [Chimura])
= Am e le tus longu lus Sinitshenk ova , 1981 = Am e le tu s s a p p o r e n s is : I ma ni shi , 1 93 2 ( non S i p h lu r us s a ppo r e n sis Ma tsu mu r a , 1904) REFERENCES. Ima nishi, 1 93 2: 52 6, Pl.31 : Fig. 1, Pl. 3 2: Figs 2 -3 ( ma le a nd fema le ima goes, as Am eletus sa pp or en sis); I ma nishi, 193 3: 6 5, Figs 3, 5 ( la rva ) ; Sinitshenk ova & T sherno va , 1 9 7 6 : 1 6 , Fi gs 1 0 - 1 7 ( ma le a nd f ema l e ima g o, ma le subimago, lar va ); Sinitshenk ova, 1 98 1: 77 , Fig. 3 ( la rva, a s A. long u lus ) ; T sher nova et a l ., 1 9 86 : 1 26 , F igs 5 7 : 1 , 2 ( ma le i m a g o ). MATERIAL EXAMINED . Among the sp ecimens list ed by Sinit she nko va & Tsher nov a [1976], the following o nes ar e rede ter mined by me as A. co stalis: RU SSIA: P RIM ORSKIY KRAY: Olginskiy Rayo n ( = Olgino Dist r.) , Furm ano vo, 15 .VI.19 72 (A. Ra snitsy n) -- 1 Im #; Kav ale rov skiy Ra yon (= Ka valero vo Distr .), Pe rev al'noy e, 29. VI.1 972 (A. P ono mar enko) -- 1 Im #, 1 Im $; riv er Kam enka, tribut ary of riv er Sup utinka , 3 .VI.19 72 (I. L eva nido va) -- 1 S #, 1 larv al exuvia e. Sy ste mat ic position of ot her sp ecies (lar vae ) is d oub tful.

257

Lava. Labru m length 0 .93- 0.9 4 of width (2 specimens m ea su re d) ( un like s h ort er in A . a lta icus an d long er in A. p arvus). Abdomin al terga and stern a w ithou t spine-like setae (un like group A. alexand rae). Tergalii I an d II with vestigial an al ribs (the same in A. formosus an d A. atratus, un like other species), small. Tergalii III-V I w ith anal rib far from anal margin (un like A. in opin atus, A. ca mtschaticus and A. altaicus). Su bimago. Wings brow nish, with crossvein s bordered by darker brow n. Imago. Wings mostly colou rless, in anterior part crossveins darken ed and bordered by brow nish. Ma le g enitals. In im ago, lateral penis lobes with apices sligh tly bent medially-dorsally; each ventral plate bears a single long denticle wh ich stretched closely parallel to the lateral lobe (u nlike all other P alaearctic species). Su bimagin al penis also h as one pair of large den ticles corresponding to im aginal ones. Egg. Flattened, in a form of ellipse-like bicon vex len s. Reticu lation: each polygonal cell with flat bottom lacking protuberance, with borders mu ch lower and n arrower th an that of roun d cells, so rou nd cells (each bearin g papilla) promin ent; round cells are small and sparse on most su rface, dense and large on one pole (the sam e in A. mon tanus and A. ced rensis). Size. Large: fore wing length 15-1 8 m m. DISTRIBUT ION . Ru ssian Far East and J apan. CO MME NTS. Th e species Ameletus lon gulus w as described as larvae only; th e description w as based on two specim ens [Sinitshenkova, 198 1]. According to the origin al description, A. lo ngu lus differs from A. costalis by longer vestiges of ribs on tergalii I, longer tergalii II an d some oth er ch aracters. Possibly, these characters vary individu ally. Th e n ame Siphlu rus sap porensis Matsumu ra, 190 4 w as wrongly applied to this species by Imanish i [193 2], while actually it is an older syn onym of Cinygm ula grand ifolia Tshern ova, 19 52 [Ish iwata, 2001 ].

Lava. Labru m length 0 .77 -0.93. of width (19 specimens measu red) (u nlike shorter in A. altaicu s sp.n. and longer in A. p arvus). Abdominal terga and sterna with spin e-like setae (u nlike other Am eletidae). Tergalii I and II w ithout anal ribs (un like A. costalis), small. Tergalii III- VI w ith anal rib far from anal margin (un like A. in opin atus, A. ca mtschaticus and A. altaicus sp.n.). Su bimago. Wings brow nish, either u nicolour, or with crossvein s n arrowly bordered by darker brow n, or w ith darker brown maculae on crossvein s. Imago. Wings n on-coloured. Ma le g enitals. In im ago, lateral penis lobes with apices m oderately ben t m edially-dorsally; ven tral plates w ith out den ticles. Egg. Ellipsoid (not flattened). Reticulation: each polygonal cell with bottom bearing a protuberance, with borders lower than borders of roun d cells, so round cells (each bearing papilla) are promin ent; round cells are small and sparse on most surface, dense and large on one pole. Size. Medium: fore w ing len gth 8-1 3 m m. DISTR IBUT ION . Moun tain s of C entral Asia. TAXONO MIC CO MPOSITION. Within the group 'alexa nd rae', there are forms which differ one from anoth er by some subimaginal an d larval ch aracters -- coloration of su bim agin al wings (uniform or with bordered crossveins), developmen t of spine-like setae on abdom inal terga, stern a and tergalii, et al. Probably, some of these forms have reprodu ctive isolation, because in habit in n eighboring biotops in the same area. Complete revision of this group is rath er difficult.

NOMINA DUBIA Ame letus procerus Bajkova, 1976 -- n omen dubium
Th e species w as described by B ajkova [19 76] as larvae an d female subim ago basing on two specimens -- one larva an d on e female subim ago with its larval exuviae. Th e subim ago with larval exuviae was design ated as holotype [Bajkova, 19 76: 586 ]. In the collection of Zoological In stitute RAS (S.Petersburg), there are deposited two specimens -- the larva an d th e larval exuviae, wh ile th e female subim ago reared from th is exuviae was not passed to th e Zoological Institu te and probably is lost. Th e larva was labeled by B ajkova as "holotype" and the larval exuviae -- as "paratype" (w hile actually th e larva is paratype and the exu viae is holotype, according to th e pu blication ). T h ese two specim en s belon g to two different species. T he larva (tru e paratype) h as tergalii II longer th an next on es [Bajkova, 197 6: Fig. 19 ] an d tergalii III- VI with anal rib on an al margin; it belongs to th e species later described as A. lab iatu s Sin itsh enkova 198 1 (see above, A. inopinatus labiatus). The larval exuviae (true holotype) has tergalii II shorter than next ones and tergalii III-VI with anal rib far from anal margin [Bajkova, 1976: Fig. 20]; it belong either to the species later described as A. cedrensis Sinitshenkova 1977, or the su bspecies described here as A. mo ntan us ro ssicus subsp.n.; these two forms are hardly distinguishable by larval characters. The original description of A. procerus was based on the both specimens, so it is wrong. Th e disagreem ent between holotype design ation and m useum labels provoked confu sion . Kluge [1 982 ], basing on the mu seu m labels, redescribed un der th e name "A. pro cerus" the species w hich actu ally represen ts A . ino pin atus lab ia tus. Later this error was revealed, and basing on the holotype design ation Kluge [199 5] synon ym ized A . p ro cerus with A. mon tanus Imanishi ,193 0. Actually, larval exuviae which represent the holotype of A. procerus, can belong either to A. cedrensis, or A. montanus rossicus sub sp.n., because the both species can be found in

II. 8. Group Amele tus alexandrae Brodsky, 1930 (incl. A. asiace ntralis Soldan, 1978)
REFERENCES. Br odsky, 19 30: 69 7, Figs 2 0-2 3 ( male a nd f ema l e i ma g o, egg ) ; S i ni ts h en k ov a & T sh e r no va , 1 9 76 : 1 3, Figs 1-2 (larva ); Soldan, 1 978 : 3 79, 38 0, Figs 1- 18 (A. alexa ndrae a nd A. as iace ntr alis; lar vae); Klu ge, 199 5: 7, Fig. 1 (larval l e c t o t yp e ). MATERIAL EXAMINED . KAZAKH STAN, UZBEKISTAN, KYRGYZSTAN and TADJIKIS TAN -- many L -S- Im, la rva e, subim ago es a nd ima goe s.


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Nikita J. Kluge
Kluge N. J. 198 2. [Ne w a nd litt le kno wn may flies of the US SR Far Ea st. Fa milie s L e pt op hle b iid ae a nd S ip hlo nur id ae (E phe mer opt era) ] // Vest nik Le ningra dskogo Univer siteta . N o.9 . P .11 2-1 16 [in Russian]. Kluge N. J. 199 5. [A cat alo gue of the t ype -sp ecimens in t he co lle ction of the Zo olo gic al Instit ute , Russian Ac ade my of Sc iences. Inse cta , E phe mer opt era ]. St. -Pe ter sburg, 52 pp . [in Russian]. Kluge N.J. 2004. The phylogenetic system of Ephemeroptera // Kluwe r Acad emic Publishe rs. 44 2 p p. Kluge N. J., St ude mann D ., Land olt P. & Gonser T. 19 95. A re cla ssific ation of Sip hlo nuro ide a ( Epheme rop ter a) // Bull. So c. Ent omo l. Suisse . Vol. 68. P. 103 -13 2. Ma can T. T. 196 1. A key to the nym phs of the Brit ish sp ecies of Ep hem ero pter a // Freshwa ter Biolo gic al Asso cia tio n, Sc ientific Pub lic ation. No .20. 6 4 p p. Sinit she nko va N.D . 1 977 . May flies of the ge nus Am eletus E at on of Pr imo rye (E phe mer opt era , Siphlonurid ae) // In: [Presnov od naya fauna za p ov ed nika "Ke d ro vay a Pa d '. Tr udy Biolo go- poc hvennog o inst ituta]. T.45 . N o.14 8. P.4 4-4 9 [in Russian]. Sinit she nko va N.D . 1 981 . [New spe cie s o f m ayflie s o f t he genus Am ele tus E at o n ( Ep he m e r o p t e ra , S ip hlo nur id a e ) fr om Sikho te- Alin // In: [Be spoz vonochnye zhyvo tny e v ekosist em ah lososev yh r ek D alne go Vosto ka ]. Vla divo st ok. P. 73-78 [in Russian]. Sinit she nko va N .D. & O.A. Tshe rno va 197 6. [New da ta on Asiat ic spec ies of Am eletus E ato n ( Epheme ro pte ra, Siphlonurid ae) ] // Vest nik Mo sko vskogo Unive rsitet a. Bio lgia. Vo l.5 . P .1-19 [in Russian]. Sinit she nko va N.D . & K. V. Var ykhano va. 19 89. A new sp ecies o f t he m a yfly ge nus Ameletus E at o n ( Ep he m e rop t era, S ip hlo nur id ae) fr om Mo ng o lia // E nt omo lo g ic he skoe Ob ozr enie. Vol.68. No .3. P. 576 -58 1. So lda n T. , 1 978 Am eletus asiacentralis sp. n. fro m Uzb ekista n, with not es on A. ale xandrae (E phe mer opt era , S iphlonuridae ) // Acta ent omo l. bohemo sl. Vo l.7 5. N o.6 . P .37 9- 3 8 2. Tsher nov a O.A. , Kluge N. J., Sinit she nko va N.D. & Belov V.V., 19 86. [Orde r E phe mer opt era ] // In: Opr ede lit el' na sekomy h D al'neg o Vo sto ka SSS R [Key to the insec ts of the US SR Fa r E ast ]. Vol.1. Le ningra d, Nauka. P. 99-142 [in Russian]. Ulmer G. 19 27. Tr ichopt ere n und E phe mer opt ere n. In: Ent. E rge b nisse S chwe d. Kam t chat ka - E xp e d . 1 92 0-1922 // Ar k. f. Zoo l. Bd. 19. Nr .8. S. 1-1 7. Zloty J. 19 96. A rev ision of the Ne arc tic Am eletus m ayflies ba sed on ad ult ma les, with de scr ipt ions o f seve n new sp ec ie s ( Ep hem er op ter a: Am et et ida e) // The Ca na dian E nt om olog ist. Vol. 12 8. P .2 93 -3 46. Zloty J. & Pritchard G. 19 97. La rva e a nd adults of Am eletus ma yflies (E phe mer opt era : Amele tid ae) fr om Alb ert a // The Ca nad ian Entom olo gist. Vo l.1 29. P. 251 -28 9.

this area and have the same larval structure. So the species name Ameletus procerus Bajkova 1976 should be regarded as nomen dubium.

Ame letus cri stat us Ba jkova, 1976 -- nom en dubium
Th e species was described by Bajkova [1 976] as male and female im agoes, subimagoes an d larvae; in the list of material examin ed, th ere are referred only 2 male im agoes (including holotype) and 2 larvae (bu t no females and subimagoes!). Attributing of all these specimens to one species was n ot groun ded. Th e types sh ould be deposited in th e Zoological In stitute of Ru ssian Academy of Sciences in S-P etersbu rg [B ajkova, 19 76: 582 ], but no one specim en w as passed to this in stitu te. Probably, the holotype an d all paratype are lost. The description does not allow to determ ine on which species it w as based; m ost p robably, th is is on e of the species described above. AC KNOWLED GEMENT. Th e work was supported by federal program " Leading Scien tific Schools", project N o. 7 130 .20 06-4.

References
Ba jko va O.Ya. 197 6. [May flies of the ge nus Am eletus E at on ( Ep he m e rop t e ra) in t he Am ur b a sin] // E nt omo log ic heskoe Ob ozre nie . Vol. 55. No .3. P. 528 -588 [in Russian]. Br ekke R. 1 965 . Bidr ag tilkunnska pen om No rge s d ogn-, stenog va rfluer (E phe mer opt era , P lec opt era , Tric hop ter a) // No rsk. E nto mol. Tidsskr . Vol. 13. No .1-2. P.1 1-1 5. Brodsky K.A. 1930. Zur Kenntnis der mittelasiatischen Ephemeropteren. I. (Imagines) // Zool. Jahrb., Syst. Bd.59. S.681-720. Im anishi K. 19 32. Ma yflies fr om Jap ane se tor rents II. Further no tes on the g enus Ameletus // Ann. Zoo l. Jap on. Vo l.1 3. P . 52 5 -5 3 2. Imanishi K. 1933. Mayflies from Japanese torrents III. Third notes on genus Ameletus with a list of the Japanese Siphlonuridae // Insecta Matsumurana. Vol.8. No.2. P.64-69. Ishiwata Shin- ichi 2 001 . A checklist of Ja panese Ep hem ero pte ra // In: Y.J . Ba e ( e d . ). The 21 st Ce nt ur y and Aquat ic Entom olo gy in East Asia. Pro c. 1st S ymp . AESE A. Kor ean So c. Aquat ic Ent omo l. Kore a. P.55-8 4. Ka ng Shin-Chang & Ya ng Chung- Tu 199 4. Thr ee new sp ecies o f the ge nus Am eletus fr om Ta iwa n ( Ep he m e rop t e ra: Siphlonurid ae) // Chine se J. E nto mol. Vol. 14. P. 261 -26 9. Kluge N. J. 197 9. [A new sp ecie s o f m ayflie s o f t he genus Am eletus Ea ton (E phe mer opt era , Siphlonurid ae) fr om Easter n Kaza khstan] // E nto mologiche sko e Oboz renie. Vo l.5 8. No .4. P. 807 -810 [in Russian].