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Ent21 3 273 306 Kluge.Pmd Russian Entomol. J. 21(3): 273306 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2012 Contribution to the knowledge of Choroterpes (Ephemeroptera: Leptophlebiidae) Ê ïîçíàíèþ Choroterpes (Ephemeroptera: Leptophlebiidae) Nikita J. Kluge Í.Þ. Êëþãå Department of Entomology, St. Petersburg State University, Universitetskaya nab., 7/9, St. Petersburg 199034, Russia. E-mail: [email protected]. Website: http://www.insecta.bio.pu.ru Êàôåäðà ýíòîìîëîãèè, áèîëîãî-ïî÷âåííûé ôàêóëüòåò, Ñ.-Ïåòåðáóðãñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò, Óíèâåðñèòåòñêàÿ íàá., 7/8, Ñ.-Ïåòåðáóðã 199034, Ðîññèÿ. KEY WORDS: systematics, Ephemeroptera, Leptophlebiidae, Atalophlebomaxillata, Choroterpes, Neochorot- erpes, Euthraulus, Monophyllus, Dilatognathus, Choroterpides, new species. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: ñèñòåìàòèêà, Ephemeroptera, Leptophlebiidae, Atalophlebomaxillata, Choroterpes, Neochoroterpes, Euthraulus, Monophyllus, Dilatognathus, Choroterpides, íîâûå âèäû. ABSTRACT. The taxon Atalophlebomaxillata is char- ÐÅÇÞÌÅ. Òàêñîí Atalophlebomaxillata õàðàêòå- acterized by presence of an apical flange on maxilla; this ðèçóåòñÿ íàëè÷èåì àïèêàëüíîé ïëàñòèíêè íà ìàê- flange is lost only in those representatives, whose maxil- ñèëëå; ýòà ïëàñòèíêà óòðà÷åíà òîëüêî ó òåõ ïðåäñòà- lae are highly specialized. Within Atalophlebomaxillata- âèòåëåé, ìàêñèëëû êîòîðûõ ñèëüíî ñïåöèàëèçèðîâà- Atalophlebolinguata, a new tribe Choroterpini tribus n. íû.  ñîñòàâå AtalophlebomaxillataAtalophlebo- (or Choroterpes/fg1) is established, to comprise Thrau- linguata âûäåëÿåòñÿ íîâàÿ òðèáà Choroterpini tribus n. lus and Choroterpes s.l. (or Choroterpes/fg2). The taxon (èëè Choroterpes/fg1), îáúåäèíÿþùàÿ Thraulus è Choroterpes/fg2 is divided into Choroterpes/fg3 (charac- Choroterpes s.l. (èëè Choroterpes/fg2). Òàêñîí terized by lanceolate median filaments on tergalii) and Choroterpes/fg2 äåëèòñÿ íà Choroterpes/fg3 (õàðàêòå- Euthraulus/g1 (characterized by arched penial arms). ðèçóþùèéñÿ ëàíöåòîâèäíûì ìåäèàëüíûì ôèëàìåí- Choroterpes/fg3 includes a widely distributed plesiomor- òîì òåðãàëèé) è Euthraulus/g1 (õàðàêòåðèçóþùèéñÿ phon Choroterpes/fg4 and a Nearctic taxon Neochorot- äóãîâèäíûìè ðóêàìè ïåíèñà). Choroterpes/fg3 âêëþ- erpes. Based on imagoes reared from larvae, new species ÷àåò øèðîêî ðàñïðîñòðàí¸ííûé ïëåçèîìîðôîí and subspecies of Choroterpes/fg4 are described Ch. Choroterpes/fg4 è íåàðêòè÷åñêèé òàêñîí Neochoro- (Ch.) picteti subcostalis ssp.n. from Caucasus (Russia), terpes. Ïî èìàãî, âûâåäåííûì èç ëè÷èíîê, îïèñûâà- Ch. (Ch.) ludmilae sp.n. from Oman and Ch. (Ch.) þòñÿ íîâûå âèäû è ïîäâèäû Choroterpes/fg4 Ch. mercatorius sp.n. from Sulawesi Island (Indonesia). The (Ch.) picteti subcostalis ssp.n. ñ Êàâêàçà (Ðîññèÿ), Ch. taxon Euthraulus/g1 has exclusively Old-World distribu- (Ch.) ludmilae sp.n. èç Îìàíà è Ch. (Ch.) mercatorius tion; it includes widely distributed plesiomorphon Eu- sp.n. ñ î. Ñóëàâåñè (Èíäîíåçèÿ). Òàêñîí Euthraulus/g1 thraulus/g2 and two small Oriental taxa Monophyllus èçâåñòåí èñêëþ÷èòåëüíî èç Ñòàðîãî Ñâåòà; îí âêëþ÷à- subgen.n. and Dilatognathus subgen.n. Based on ima- åò øèðîêî ðàñïðîñòðàí¸ííûé ïëåçèîìîðôîí Euthraulus/ goes reared from larvae, a new species of Euthraulus/g2 g2 è äâà ìàëåíüêèõ îðèåíòàëüíûõ òàêñîíà is described Ch. (Euthraulus) caucasicus sp.n., and Monophyllus subgen.n. è Dilatognathus subgen.n. Ïî additional descriptions to Ch. (E.) altioculus, Ch. (E.) èìàãî, âûâåäåííûì èç ëè÷èíîê, îïèñûâàåòñÿ íîâûé sumbarensis and Ch. (E.) signatus are given. The taxon âèä Euthraulus/g2 Ch. (Euthraulus) caucasicus sp.n., Monophyllus includes a single species Ch. (Monophyl- à òàêæå äàþòñÿ äîïîëíèòåëüíûå îïèñàíèÿ Ch. (E.) lus) monophyllus sp.n., described as mature larvae from altioculus, Ch. (E.) sumbarensis è Ch. (E.) signatus. Hainan Island (China). Larvae of the taxon Dilatog- Òàêñîí Monophyllus âêëþ÷àåò åäèíñòâåííûé âèä Ch. nathus are known under the generic name Choroterpides, (Monophyllus) monophyllus sp.n., îïèñàííûé ïî çðå- whose type species (exiguus Eaton, 1884 [Thraulus]) is ëûì ëè÷èíêàì ñ î. Õàéíàíü (Êèòàé). Ëè÷èíêè Dilato- known only as imago and probably belongs to Euthrau- gnathus èçâåñòíû ïîä ðîäîâûì íàçâàíèåì Choro- lus/g2. The type species of Dilatognathus is Choroterpes terpides, òèïîâîé âèä êîòîðîãî (exiguus Eaton, 1884 (Dilatognathus) cataractae sp.n., described as imagoes [Thraulus]) èçâåñòåí òîëüêî ïî èìàãî è, âåðîÿòíî, reared from larvae in Lombok Island (Indonesia). Similar îòíîñèòñÿ ê Euthraulus/g2. Òèïîâûì âèäîì Dilato- modifications in mouthparts structure independently gnathus subgen.n. ÿâëÿåòñÿ Choroterpes (Dilato- evolved in non-related taxa Dilatognathus, Notophlebia, gnathus) cataractae sp.n., îïèñàííûé ïî èìàãî, âûâå- Hermanellognatha, Hagenulus/fg3 and Ulmeritus/g2. äåííûì èç ëè÷èíîê íà î. Ëîìáîê (Èíäîíåçèÿ). Ñõîä- 274 Nikita J. Kluge íûå ïðåîáðàçîâàíèÿ â ñòðîåíèè ðîòîâûõ ÷àñòåé íå- Petersburg, Russia), but is temporarily located in the çàâèñèìî ïðîèçîøëè ó íåðîäñòâåííûõ òàêñîíîâ Department of Entomology of Saint Petersburg State Dilatognathus, Notophlebia, Hermanellognatha, Hage- University. In the lists of material examined, the follow- nulus/fg3 è Ulmeritus/g2. ing arbitrary signs are used: L larva; S subimago; I imago; L-S-I# male imago reared from larva, Introduction with larval and subimaginal exuviae; L-S-I$ female imago reared from larva, with larval and subimaginal This paper continues revision of mayfly system based exuviae; L-S# male subimago reared from larva, on the principles of cladoendesis [Kluge & Novikova, with subimaginal exuviae; L/S# male subimago 2011; Kluge, 1912] with usage of the dual nomenclature extracted from mature larva. system [Kluge, 2010]. Based on these principles, in- 1. ATALOPHLEBOMAXILLATA, or Atalophlebia/fg4 stead of the formal subfamily Atalophlebiinae, a series of subsequently subordinated rank-free taxa have been Systematic position: Leptophlebia/fg1Atalophleboadentata established: Atalophleboadentata (or Atalophlebia/fg1) AtalophlebopectinataAtalophleboculataAtalophlebomaxillata. Circumscriptional name: Atalophlebomaxillata Kluge, 2009. > Atalophlebopectinata (or Atalophlebia/fg2) > Hierarchical typified name: Atalophlebia/fg4 (sine Terpides; Atalophleboculata (or Atalophlebia/fg3) > Atalophle- incl. Castanophlebia). bomaxillata (or Atalophlebia/fg4) > Atalophlebolin- Possible rank-based name: guata (or Atalophlebia/fg5) [Kluge, 2004, 2008, 2009]. subfamily Atalophlebiinae s.l. The phylogenetic relationships within the taxon Autapomorphies. (1) Maxilla has an apical flange, which represent a Atalophlebolinguata remain poorly known. Within continuation of its ventral wall to apical direction and sepa- Atalophlebolinguata, a subordinated holophyletic tax- rates the apical field of long filtering setae (characteristic for on Hagenulus/fg1 has been established [Kluge, 2008]. Leptophlebia/fg1) from the apical-ventral row of pectinate In the present paper, based on new material, the setae (also characteristic for Leptophlebia/fg1); thanks to the taxon Atalophlebomaxillata is redefined and a new apical flange, the apical field of long filtering setae is shifted holophyletic taxon Choroterpes/fg1 within Atalophle- to the dorsal side of maxilla and its ventral margin locates far bolinguata is established. Classification of the taxa dis- from the apical-ventral row of pectinate setae (Fig. 3). The cussed in this paper is the following: apical flange reaches inner margin of maxilla, and laterally terminates fare from outer margin of maxilla (Fig. 2). The 1. Atalophlebomaxillata apical flange is very thin and translucent, so it is poorly 1.1. Atalophlebolinguata visible on background of dense apical filtering setae. In my 1.1.1. Choroterpes/fg1 (incl. Thraulus) previous paper maxilla of Farrodes bimaculatus Peters & 1.1.1.1. Choroterpes/fg2 (incl. Euthraulus) Alayo, 1971 was shown wrongly, without the apical flange 1.1.1.1.1. Choroterpes/fg3 (incl. Neochoroterpes) [Kluge, 1994: Fig.4]; actually it has the apical flange similar 1.1.1.1.1.1. Plesiomorphon Choroterpes/fg4 to that in Fig. 2. 1.1.1.1.1.1-1. Choroterpes/fg4 picteti [Choroterpes] Among the taxa examined, the apical flange is reduced 1.1.1.1.1.1-1.1. Choroterpes/fg4 p. picteti [Choroterpes] only in taxa with highly specialized maxillae of the «Dilatog- nathus-like type» Hermanellognatha [Kluge, 2008: Fig. 1.1.1.1.1.1-1.2. Choroterpes/fg4 p. subcostalis ssp.n. [Choroterpes] 37], Notophlebia Peters & Edmunds, 1970, Hagenulus mor- 1.1.1.1.1.1-2. Choroterpes/fg4 nigrescens [Choroterpes] risonae Peters & Alayo, 1971 (but not Hagenulus caligatus 1.1.1.1.1.1-3. Choroterpes/fg4 ludmilae sp.n. [Choroterpes] Eaton, 1882) and last instar larva of Dilatognathus subgen.n. 1.1.1.1.1.1-4. Choroterpes/fg4 mercatorius sp.n. [Choroterpes] (Fig. 129) (but not its younger larvae Fig. 132). 1.1.1.1.1.2. Neochoroterpes/g(1), or Neochoroterpes Unlike Atalophlebomaxillata, in other Leptophlebia/fg1 1.1.1.1.1.2-1. Neochoroterpes/g(1) oklahoma [Choroterpes] (i.e., Terpides/fg1, Habrophlebia/fg1, Calliarcys/fg(1) and 1.1.1.1.2. Euthraulus/g1 (incl. Monophyllus, Dilatognathus) the plesiomorphon Leptophlebia/fg2) the apical flange is 1.1.1.1.2.1. Plesiomorphon Euthraulus/g2 absent, the apical field of long filtering setae is located at 1.1.1.1.2.1-1. Euthraulus/g2 altioculus [Choroterpes] most on ventral side of maxilla, and its ventral margin is 1.1.1.1.2.1-2. Euthraulus/g2 sumbarensis [Choroterpes] located close to the apical-ventral row of pectinate setae (Fig. 1) [Kluge, 2009: Fig.1]. 1.1.1.1.2.1-3. Euthraulus/g2 caucasicus sp.n. [Choroterpes] (2) Distal
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