Russian Entomol. J. 20(1): 156 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2011 Systematics of the mayfly taxon Acentrella (Ephemeroptera: Baetidae), with description of new Asian and African species Ñèñòåìàòèêà ïîäåíîê òàêñîíà Acentrella (Ephemeroptera: Baetidae) ñ îïèñàíèåì íîâûõ âèäîâ èç Àçèè è Àôðèêè Nikita J. Kluge1 & Eugenia A. Novikova2 Í.Þ. Êëþãå1, Å.À. Íîâèêîâà2 1 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 1 Êàôåäðà ýíòîìîëîãèè, áèîëîãî-ïî÷âåííûé ôàêóëüòåò, Ñ.-Ïåòåðáóðãñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò, Óíèâåðñèòåòñêàÿ íàá., 7/8, Ñ.-Ïåòåðáóðã 199034, Ðîññèÿ. 2 Department of Ichthyology and Hydrobiology, St. Petersburg State University, Vasilievskiy Ostrov, 16th line, 29, St. Petersburg 199034, Russia. 2 Êàôåäðà èõòèîëîãèè è ãèäðîáèîëîãèè, áèîëîãî-ïî÷âåííûé ôàêóëüòåò, Ñ.-Ïåòåðáóðãñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò, 16 ëèíèÿ ÂÎ, 29, Ñ.-Ïåòåðáóðã 199034, Ðîññèÿ. KEY WORDS: systematics, morphology, Ephemeroptera, cladoendesis, microlepides, unistyliger, penial bridge, gonovectis, Protopatellata, Anteropatellata, Baetovectata, Baetungulata, Baetofemorata, Baetosternata, Acentrellini, Acentrella, Jubabaetis, Tanzaniops, Liebebiella, Platybaetis, new species. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: ñèñòåìàòèêà, ìîðôîëîãèÿ, Ephemeroptera, êëàäîýíäåçèñ, ìèêðîëåïèäû, óíè- ñòèëèãåð, ãîíîâåêòèñ, Protopatellata, Anteropatellata, Baetovectata, Baetungulata, Baetofemorata, Baetosterna- ta, Acentrellini, Acentrella, Jubabaetis, Tanzaniops, Liebebiella, Platybaetis, íîâûå âèäû. ABSTRACT. This paper continues publications on includes 2 species inexpectatum [Pseudocloeon] and phylogenetic analysis of Ephemeroptera according to the gnom [Pseudocloeon]; their larvae, subimagoes, ima- approach, which is named here cladoendesis. A new goes and eggs are described. Group sibiricum includes 3 term microlepides is suggested for peculiar cuticular species sibiricum [Pseudocloeon], diptera sp.n. [Acen- texture. Autapomorphies of Liberevenata in genital struc- trella] (new species from Siberia and Russian Far East) ture are described, new terms unistyliger, penial bridge and scabriventris sp.n. [Acentrella] (new species from and gonovectis are introduced. Autapomorphy of An- Kyrgyzstan and Tajikistan); their larvae, subimagoes, teropatellata in tibial structure is re-defined, and a new imagoes and eggs are described. African and Oriental circumscriptional taxon name Protopatellata taxon n. is representatives of Acentrella/fg1 constitute a holophylet- established. In Anteropatellata, the following new subse- ic taxon Jubabaetis/g1 characterized by an autapomor- quently subordinated holophyletic taxa are established phy in structure of left mola. Jubabaetis/g1 is divided to and characterized: Baetovectata taxon n., Baetungulata African taxon Tanzaniops/g(1) and Oriental taxon Juba- taxon n., Baetofemorata taxon n. and Baetosternata baetis/g2. A new species lunamontana sp.n. [Acentrella taxon n. A new typified name Acentrellini tribus n. is (Tanzaniops)] is described as larvae, subimagoes and introduced, to form a hierarchical name Acentrella/fg1 imagoes from Uganda. Holophyletic taxon Jubabaetis/g2 for a taxon belonging to Baetofemorata-non-Baetoster- is divided into a plesiomorphon Liebebiella/g(1) and nata. System of Acentrella/fg1 is revised. All Holarctic holophyletic taxa Jubabaetis/g3 and Platybaetis/g(1). representatives of Acentrella/fg1 are temporarily placed Three species of Liebebiella/g(1) are described as larvae, to a plesiomorphon Acentrella/fg2. Key to larvae and subimagoes and imagoes: verum [Pseudocloeon] (= deig- male imagoes of Palearctic species of Acentrella/fg2 is ma [Liebebiella], syn.n.), bispinosa sp.n. [Acentrella given; they are divided into four species groups. Group (Liebebiella)] and cylindroculata sp.n. [Acentrella (Lie- lapponica is a plesiomorphon, characterized by presence bebiella)] (new species from Sulawesi). Two species of of hind wings; larvae, subimagoes, imagoes and eggs are Platybaetis/g(1) from Sulawesi are described as larvae, described for lapponica [Acentrella], charadra [Acen- subimagoes and imagoes: mamasae sp.n. [Platybaetis] trella] and chantauensis [Baetis]; feropagus [Acentrel- and wallacei [Platybaetis]. la], known as larvae only, is redescribed basing on mate- rial from far north-east of Russia; comments on joosti ÐÅÇÞÌÅ. Äàííàÿ ñòàòüÿ ïðîäîëæàåò ñåðèþ ïóá- [Baetis] are given. Group fenestratum includes a single ëèêàöèé ïî àíàëèçó ôèëîãåíèè ïîä¸íîê â ñîîòâåò- species fenestratum [Pseudocloeon]; larvae, subimagoes, ñòâèè ñ ïîäõîäîì, êîòîðûé íàçâàí çäåñü êëàäîýíäå- imagoes and eggs are described. Group inexpectatum çèñ. Íîâûé òåðìèí ìèêðîëåïèäû ïðåäëîæåí äëÿ 2 N.J. Kluge, Eu.A. Novikova îñîáîãî ðåëüåôà êóòèêóëû. Îïèñûâàþòñÿ àóòàïîìîð- Ephemeroptera [Kluge, 2004]. Updated electronic ver- ôèè Liberevenata â ñòðîåíèè ãåíèòàëèé è ââîäÿòñÿ sion of this monograph is available from íîâûå òåðìèíû óíèñòèëèãåð, ìîñò ïåíèñà è ãî- www.insecta.bio.pu.ru. The method of phylogenetic íîâåêòèñ. Ïåðåîïèñàíà àóòàïîìîðôèÿ Anteropatellata analysis used there can be named cladoendesis i.e., â ñòðîåíèè ãîëåíè è óñòàíîâëåí íîâûé òàêñîí branch-coupling (from Greek êëáäïò + åí-äåóéò). Protopatellata taxon n.  ñîñòàâå Anteropatellata óñòà- Cladoendesis is based on a specially elaborated rational íîâëåíû è îõàðàêòåðèçîâàíû íîâûå ïîñëåäîâàòåëü- text layout and a powerful, rank-free, dual nomencla- íî ñîïîä÷èí¸ííûå òàêñîíû Baetovectata taxon n., ture system [Kluge, 2010]. This allows to arrange all Baetungulata taxon n., Baetofemorata taxon n. è characters among taxa, make clear links between char- Baetosternata taxon n. Ââîäèòñÿ íîâîå òèïèôèöèðî- acters of subordinated and higher taxa, clarify phyloge- âàííîå íàçâàíèå Acentrellini tribus n., ÷òîáû îáðàçî- netic significance of characters, adequately use in phy- âàòü èåðàðõè÷åñêîå íàçâàíèå Acentrella/fg1 äëÿ òàê- logenetic analysis all know apomorphies, and find ways ñîíà, îòíîñÿùåãîñÿ ê Baetofemorata-non-Baetosternata. to discover new apomorphies [Kluge, 2000; 2004]. Ïåðåñìàòðèâàåòñÿ ñèñòåìà Acentrella/fg1. Âñå ãîëàð- Cladoendesis is opposed to a non-scientific approach of êòè÷åñêèå ïðåäñòàâèòåëè Acentrella/fg1 âðåìåííî counting characters and calculating parsimony (which ïîìåùàþòñÿ â ïëåçèîìîðôîí Acentrella/fg2. Äàþòñÿ is not compatible with any modern evolutionary theo- êëþ÷è äëÿ îïðåäåëåíèÿ ëè÷èíîê è ñàìöîâ èìàãî ry). The phylogenetic system of Ephemeroptera gives ïàëåàðêòè÷åñêèõ âèäîâ Acentrella/fg2, êîòîðûå äå- detailed cladoendetic system of all mayflies except for ëÿòñÿ íà 4 ãðóïïû âèäîâ. Ãðóïïà lapponica ÿâëÿåòñÿ Leptophlebiidae and Baetidae. The present paper starts ïëåçèîìîðôîíîì, õàðàêòåðèçóþùèìñÿ íàëè÷èåì çàä- a series of publications on cladoendesis of Baetidae. íèõ êðûëüåâ; îïèñûâàþòñÿ ëè÷èíêè, ñóáèìàãî, èìà- For most species, described in this paper, imagoes ãî è ÿéöà lapponica [Acentrella], charadra [Acentrella] were reared from larvae in field conditions during trips è chantauensis [Baetis]; feropagus [Acentrella], èçâåñò- to various regions of Russian Federation (Urals, Sibe- íûé òîëüêî ïî ëè÷èíêàì, ïåðåîïèñûâàåòñÿ ïî ìàòå- ria, Far East), Kyrgyzstan, Tajikistan, Mongolia, Indo- ðèàëó ñ êðàéíåãî ñåâåðî-âîñòîêà Ðîññèè; äàíû çàìå- nesia (Java, Sulawesi) and Uganda. All material, includ- ÷àíèÿ î joosti [Baetis]. Ãðóïïà fenestratum âêëþ÷àåò ing holotypes and paratypes of new species, is perma- åäèíñòâåííûé âèä fenestratum [Pseudocloeon]; îïè- nently deposited in the Zoological Institute of Russian ñûâàþòñÿ åãî ëè÷èíêè, ñóáèìàãî, èìàãî è ÿéöà. Ãðóï- Academy of Sciences (Saint Petersburg, Russia); tem- ïà inexpectatum âêëþ÷àåò 2 âèäà inexpectatum porarily locates in the Department of Entomology of [Pseudocloeon] è gnom [Pseudocloeon]; îïèñûâàþòñÿ Saint Petersburg State University. In the lists of material èõ ëè÷èíêè, ñóáèìàãî, èìàãî è ÿéöà. Ãðóïïà sibiricum examined, the following arbitrary signs are used: L âêëþ÷àåò 3 âèäà sibiricum [Pseudocloeon], diptera larva; S subimago; I imago; L-S-I# male sp.n. [Acentrella] (íîâûé âèä èç Ñèáèðè è Äàëüíåãî imago reared from larva, with larval and subimaginal Âîñòîêà) è scabriventris sp.n. [Acentrella] (íîâûé âèä exuviae; L-S-I$ female imago reared from larva, èç Êèðãèçèè è Òàäæèêèñòàíà); îïèñûâàþòñÿ èõ ëè- with larval and subimaginal exuviae; L-S# male ÷èíêè, ñóáèìàãî, èìàãî è ÿéöà. Àôðèêàíñêèå è îðè- subimago reared from larva, with subimaginal exuviae; åíòàëüíûå ïðåäñòàâèòåëè Acentrella/fg1 îáðàçóþò L/S# male subimago extracted from mature larva. ãîëîôèëåòè÷åñêèé òàêñîí Jubabaetis/g1, õàðàêòåðè- Before description of taxa, it is necessary to give two çóþùèéñÿ àóòàïîìîðôèåé â ñòðîåíèè ëåâîé ìîëû. comments: (1) on status of the old generic name Pseudoc- Jubabaetis/g1 äåëèòñÿ íà àôðèêàíñêèé òàêñîí loeon and (2) on a new term for description of color patterns. Tanzaniops/g(1) è îðèåíòàëüíûé òàêñîí Jubabaetis/ g2. Ïî ëè÷èíêàì, ñóáèìàãî è èìàãî èç Óãàíäû îïè- A. General remarks ñûâàåòñÿ íîâûé âèä lunamontana sp.n. [Acentrella (Tanzaniops)]. Ãîëîôèëåòè÷åñêèé òàêñîí Jubabaetis/ A.1. Status of the generic name Pseudocloeon g2 äåëèòñÿ íà ïëåçèîìîðôîí Liebebiella/g(1) è ãîëî- Most species, which are here attributed to Acentrel- ôèëåòè÷åñêèå òàêñîíû Jubabaetis/g3 è Platybaetis/g(1). la/fg1, have been originally described in the genus Ïî ëè÷èíêàì, ñóáèìàãî è èìàãî îïèñûâàþòñÿ òðè Pseudocloeon. The generic name Pseudocloeon Kla- íîâûõ âèäà Liebebiella/g(1): verum [Pseudocloeon] (= palek, 1905 is older than Acentrella