8 Zoological Institute, St.Petersburg, 1997

New subgenera of Holarctic (Ephcmeroptera: , , Ephemera:llidae)

N.Ju Kluge

Kluge, N.Ju. 1997. New subgenera of Holarcti~:mayflies (Ephemeroptera: Heptagenii- dae, Leptophlebiidae, ). Zoosystematicc~Rossica, 5(2), 1996: 233-235.

Three new subgenera are described: (1) Caucasiron subgen. n. (type caucasica Tshernova, 1938) of the s. I. ~(Heptageniidae),which includes several Caucasian and Middle Asian species formerly placed in the subgenus (or ge- nus) Iron; (2) Neoleptophlebia subgen. n. (type species chocolata Imanishi, 1937) of the genus s. 1. (Leptophlebiidae), which includes sev- eral Asian and North American species formerly placed in the subgenus (or genus) Paraleptophlebia; (3) monotypic subgenus Arnurella subgen. n. (type species Ephe- merella gracilis Tshernova, 1952) of the genus Ephemerella s. 1. (Ephemerellidae).

N. Ju. Kluge, Department of Entomology, Biological fizculty, St.Petersburg State Uni- versity, Universiterskaya nab. 7, St.Petersburg 199034, ,?ussia.

Family HEPTAGENIIDAE Traver, 1935). One more subgenus of Epeorus s. 1. with a suction disc is described here. Genus Epeorus Eaton, 1881 (s. 1.) Subgenus C:aucasironsubgen. n. Type species Epeorus torrentium Eaton, 1881. The genus Epeorus is accepted here in the Type species Cinykma caucasica Tshernova, 1938. broad sense (see Kluge, 1988, 1993). Larvae Diagnosis. Lama. Hairs on anterior mar- of the subgenera Epeorus s. str. and Belovius gin of head directed dorso-medially (as in Tshernova, 198 1 have tergaliae not forming Ironopsis, in contrast to other Epeorus s. l., complete suction discs. The species whose where these hairs are directed forward). Ter- larvae have suction discs were initially galiae form coaiplete suction disc (as in placed by Eaton in the genus Iron Eaton, Ironopsis and many species of Iron s. str., in 1885; the name "Iron" means "a dissembler", contrast to Epeorus s. str. and Belovius); ter- that is associated with the fact that larvae of galiae of pair I st:rongly expanded anteriorly this genus have the same structure of suction and contiguous under thorax; tergaliae of disc as in the genus Rhilhrogena, but in all pair VII with loingitudinal fold which per- other respects resemble Epeorus (Eaton, mits them to turn under apex of abdomen. 1883- 1888). A suction disc of the same s truc- Tergaliae of pairs I-VII (or at least anterior- ture appears in several genera of Ephe- most of them) with distinct projection on an- meroptera, namely in and terior-dorsal side of anterior margin (Sinit- Epeorus s. 1. (Heptageniidae) and in Delea- shenkova, 1976: IFigs 31, 32; Braasch, 1978: ridium and Lepeorus (Leptophlebiidae). As Fig. 10; 19'79: Figs lp, lq, 2i, 2j; Braasch & species of Epeorus s. 1. with suction discs Soldan, 1979: Fig. 6) (UI contrast to all other have no other common characters separat- Epeorus s. 1. and Bphemeroptera in general). ing them from other Epeorus s. I., mono- Imago. Penis lobes with a pair of titillators phyly of this group is not proved. At pre- (in contrast to Ironopsis), without a pair of sent, the representatives of Epeorus s. 1. with dorsolateral denticles (in contrast to Iron s. suction discs are placed in two separate sub- str.) (Tshernova, 1938: Fig. 5; Sinitshenk- genera (or genera): Iron s. str. (with type spe- ova, 1976: Figs 1-3; Braasch, 1979: Figs le, cies I. longimanus Eaton, 1885) and Ironopsis If, 2d, 2f, 3c, 4e, 4c). Traver, 1935 (with type species Iron grandis Distribulion. Caucasus and Middle Asia. 2 34 N. Ju. Huge: New subgenera ofmuyj2ies . 7,OOSYST. ROSSICA Vol. 5 Species examined. E. (C.} c~lucasicus Species examined L. (P.) cincta (Retzius, (Tshernova, 1938) (originally in Cinygina): 1783): imagos reared from larvae; L. (P.) lectotype and paralectotypes (Kluge, 1995) longilobata Tshernova, 1928: imagos - lecto- and imagos reared from larvae; E. (C.) zno- type and paralectotypes (Kluge, 1995); L. jkoi (Tshernova, 1938) (originally Ln Iron): (P.) strandii Eaton, 1901 [= L. (P.) lunata lectotype (Kluge, 1995) and imagos reared Tshernova, 19521: imagos reared from lar- from larvae; E. (C.) alpesrris (Braasch, vae; L. (P.) submarginata (Stephens, 1835): 1979) (originally in Iron): lmagov reared imagos reared from larvae; L. (P.) werneri from larvae; E. (C.) magnus (Braasch, 1978) (Ulmer, 19 19): imagos reared from larvae; L. (originally in Iron): imagos reared fiom lar- (P.) guttara McDunnough, 1924: imagos vae; E. (C.) soldani (Braasch, 197!3) (origi- reared from larvae; L. (P.) packii Needham, nally in Iron): imagos reared From larvae; E. 1927: larvae. (C.) guttalus (Braasch & Soldan. 1979): imagos reared from larvae. Subgenus Neoleptophlebia subgen. n.

Family LEPTOPHLEBIIDAE Type species Paraleprophlebia chocolata Imanishi, 1937. Genus Leptophlebia Westwood, 1840 (s. 1.) Diagnosis. . 'Tergaliae furcate far from their base; tergalial tracheae usually Type species Ephemeru vespertine: I.~nnac:us,1958. with additional branches, especially proxi- Here this genus is accepted in the broad mally from the furcation. Terminal segment sense and includes the subgenus Pvalepro- of labial palpi widest near its base. phlebia Lestage, 1917. At present the subge- Male imago. Penis lobes without ventral nus (or genus) Paraleptophlebia is charac- appendages arising at apex and directed ven- terized only by the absence of apomorphies trally (other appendages may be present). peculiar to the subgenus (or genus) Lepto- Relarionship. The shape of larval tergaliae phlebiu s. str.: branches of tergaliae 11-VII in and palpi in Neoleprophlebia resemble those Paraleprophlebia are not widened, imaginal of , Calliarcys and Habro- penis of various structure, but not as in Lep- phlebiinae and probably are synapomor- tophlebia s. str. (Peters & Etlmunds, 1970). phies of these taxa, indicating paraphyletic Therefore Paraleprophlebia s. 1. is regarded nature of the genus Leptophlebia s. 1. as a paraphyletic taxon. To make it more Disrribulion. Eastern Palaearctic, Oriental natural, the former subgenus Paralqtophk- Region, Nearctic. bia s. 1. is divided here into two subgenera: Species examined. L. (N.) chocolata Paraleptophlebia s. str. and Neoleprophlebia (Imanishi, 1937): imagos reared from larvae; subgen. n. L. (N.) vladivosrokica (Kluge, 1982): imagos reared from larvae; L. (N.') mollis (Hagen, Subgenus Paraleptophlcbia Lestage, 191 7 (s. str.) 186 1): imagos reared from larvae; L. (N.) adopliva McDunnough, 1929: larvae. Type species Eplren~eracincta Retzius, 1'183. Diagnosis. Larva. Tergaliae with two lobes Fa mil y EPHEMERELLIDAE separated nearly up to base (in contrast to Neoleptophlebia), lobes of all tergaliae slen- Cienus Ephemerella Walsh, 1862 der (in contrast to Lepfophlebia s. str.); ter- galial tracheae without additional t~ranches. Type species Ephemerella excruciam Walsh, 1862. Terminal segment of labial palpi widest near Here this genus is accepted in wide sense: its middle. it corresponds to the subtribe Ephemerellae Male imago. Penis with a pair of ventral sensu Allen, 1984 and includes subgenera appendages arising from apex of each penis Drunella Needham, 1 905, Caudalella Ed- lobe and directed proximally. munds, 1959, CincticostelZa Allen, 197 1, Ac- Relarionship. The presence of peculiar ven- erella Allen, 197 1, Serratella Edmunds, 1959, tral appendages of the penis in male imago Torleya Lestage, 19 17, Uracanthella Belov, and the deep separation of tergaliae in larvae 1979, Caurinella Allen, 1984, Teloganopsis are probably synapomorphies of Fi~ralepto- Ulmer, 1939, Crinirella Allen & Edmunds, phlebia s. str, and Leptophlebia s. str. indicat- 1963 (some authors regard these taxa as ing that these two taxa form a holophyletic separate genera). The genus Ephemerella s. 1. group separate from hreoleptoph1ebii.r. can be separated from other Ephemerellidae Disrriburion. Holarctic. by the absence of rudiments of tergaliae on ZOOSYST. ROSSICA Vol. 5 N.Ju. Kluge: Arew subgenera of madlies 235 the abdominal segment I and presence of aus dem Kaukas~~s.Entomol. Nachr., 22(5): 65- tergaliae only on segments 111-VII. 70. Braasch, D. 1979. Beitrag zur Kenntnis der Gattung Iron Eaton im Kaukasus (UdSSR) (111) (Ephe- Subgenus mur re kt subgen. n. meroptera, Heptageniidae). Reichenhachia, 17(33): 283-394. Type species Ephenlerelia gracilis Tshernova, 1952. Braasch, D. & Soldan, T. 1979. Neue Heptageniidae Diagnosis. Larva. Body slender (in con- aus Asien (E~~hemeroptera). Reichenbachia, trast to Torleya and some others). Maxillae 17(31): 261-272. biting, not filtering (in contrast to Uracan- Eaton, A.E. 1883- l Fl88. A revisional monograph of thella and Citrcticosfella). Fore legs not recent or mayflies. Trans. Linn. Soc. grasping (in contrast to Drunella). Pronotum Lond. 3(2): 1-352,65 pl. with straight anterior margin (in contrast to Kluge, N.Ju. 1988. Itevision of genera of the family Heptageniidae (Ephemeroptera). I. Diagnoses of Cincticostella). Mesonotum without lateral tribes, genera artd subgenera of the subfamily projections (in contrast to AcereNa). Ab- Heptageniinae. Entomol. Obozr., 67(2): 291 -31 3. dominal terga with unpaired dorsal tubercles (In Russian). (in contrast to all other Ephemerella s. I., Kluge, N.Ju. 1993. Revision of genera of the family whose dorsal tubercles are paired or absent). Heptageniidae (ISphemeroptera). 11. Phylogeny. All tergaliae of abdominal segments 111-VI Entomol. Obozr., 72(1): 39-54. (In Russian). with bifurcate ventral lamella (as in Ser- Huge, N.Ju. 1995. 11 calalogue of the fype-specimens rarella, Torleya and Uracanthella; in contrast in the coliection of the Zoological Institute. Rus- sian Academy oj' Sciences. Inrecta, Ephemerop- to Ephemerella s. str., Drunella, Caudatella tera. 52 p. St.Petersburg. (In Russian). and Cincricostella whose tergaliae of seg- Peters, W.L. & Elmunds, G.F., Jr. 1970. Revision ment VI have unfurcate ventral lamella). of the generic classification of the Eastern Nemi- Distribution. Asian Far East (rivers Amur, sphere Leptophlebiidae. Pacific , 12: 157- Ussuri and Sungari). 240. Species examined. E. (A.) gracilis 'Tsher- Sinitshenkova, N.D. 1976. Mayflies of the genus nova, 1952: lectotype and paralectotypes lron Eaton (Ephemeroptera, Heptageniidae) in the fauna of the Caucasus. Entomol. Obozr., (Kluge, 1995); larvae and subimago ex- 54(4): 853.862 (In Russian). tracted from larva. lmagos of this species are Tshernova, O.A. 1938. Zur Kenntnis der Ephe- uriknown. meropleren Ost-Transkaukasiens. Trudy 2001. inst. Azerb. Fil. .4kud. Nauk SSSR, 7(42): 55-64. References (In Kussi;an).

Braasch, 1). 1978. Epeorus znojkoi Tshern. und lron Received 30 November 1995 magnus n. sp. (Heptageniidae, Ephemeroptera)