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Phylogeny and Higher Classification of Ephemeroptera

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Phylogeny and Higher Classification of Ephemeroptera Q Zoological Institute, St.Petersburg, 1998 Phylogeny and higher classification of Ephemeroptera N.Ju. Kluge Kluge, N.Ju. 1998. Phylogeny and higher classification of Ephemeroptera. Zoosj~s- tematica Rossica, 7(2): 255-269. The order Ephemeroptera is divlded into Permoplectoptera and Euplectoptera; Eu- plectoptera are divided into two holophyletic taxa - Posteritorna and Anteritorna; An- teritorna are divided into Tridentiseta (which is possibly a paraphyletic group) and a holophyletic taxon Bidentiseta; Bidentiseta are divided into Branchitergaliae and Fur- catergaliae. Characters of these taxa are described and discussed, together with charac- ters of higher taxa in Ephemeroptera suggested by other authors. N.Ju. Kluge, Department of Enromology, Biological Faculty, St.Petersburg State Uni- versity, Universitetskaya nab. 7, Sr.Petersburg 199034, Russia. INTRODUCTION were united with some other taxa into a sub- order which was regarded as holophyletic This paper represents a report made at the (Pannota, Furcatergalia, or Rectracheata). VII International Conference on Epheme- But earlier an opposite idea was expressed, roptera in 1992 in Orono, Mine, USA (Klu- according to which Baetiscidae and Prosop- ge, 1992a), and the manuscript was submit- istomatidae separated early from other ted to the Proceedings of the Conference. Ephemeroptera and inherited their unusual Because of a confusion, the paper was not fore wing venation with unforked CuA from published in the Proceedings which appeared the Paleozoic Misthodotidae (belonging to in 1995. In spite of that, the paper was cited the extinct suborder Permoplectoptera), as a published one (Kluge et al, 1995), and while other recent mayflies originated from the names of new taxa introduced in this pa- another Palaeozoic family of the suborder per were used (Kluge, 1992b, 1993a, 1993b; Permoplectoptera -- Protereismatidae (Ed- Kluge et al., 1995). Here this paper is given munds & Traver, 1954a; etc.). with some later corrections. In the present paper, a new version of McCafferty & Edmunds (I 979) divided all mayfly phylogeny and a new classification of recent Ephemeroptera into two suborders, Ephemeroptera are suggested. It has been al- Schistonota and Pannota. Kluge (1 989) sug- ready published briefly in some papers gested another division of recent Ephe- (Kluge, 1992a, I992b, 1993b; Kluge et al., meroptera into two suborders, Costatergalia 1995). This classification is given below. and Furcatergalia. McCafferty (1 99 1) di- vided recent Ephemeroptera into three sub- TA4XONOMICPART orders: Pisciforma, Setisura and Rectra- cheata. In all these three classifications, one of the suborders (Schistonota, Costatergalia and Pisciforma) is wittingly paraphyletic, while the others (Pannota, Furcatergalia, The order is divided into (1) Permoplec- Setisura and Rectracheata) were regarded to toptera and (2) Euplectoptera. be holophyletic. In these three successive classifications, the volume of the para- 1 . Permoplectoptera Tillyard, 1932 phyletic taxon decreased, while the volume of holophyletic taxa increased. This corre- An extinct (Permian - Triassic) taxon, sponds to the general direction in which all characterized by homonornous wings and classification of organisms changes. In each of the three classifications, the some other plesiomorphies. A paraphyletic families Baetiscidae and Prosopistomatidae taxon ancestral to Euplectoptera. 256 N.Ju. Muge;Classificar,ion oJEphemeroptercr ZOOSYST. ROSSICA Vol. 7 2. Euplectoptera Tilly ard, 1932 Anteritorna are divided into (2.2.1) Tri- dentiseta and (2.2.2) Bidentiseta. Characterized by heteronomous wings: hind wing not exceeds 112 of fore wing 2.2.1. Tridentiseta Kluge et al., 1995 length; in flight, anterior margin of hind wing coupled with posterior margin of fore This group corresponds in its content to: wing. This may be an autapomorphy or a re- - Pisciforma sensu Kluge et al., 1995: 105 sult of convergence (see below). Correspond- (non Pisciforma McCafferty, 1991); ingly, this taxon may be holophyletic or - Tridentiseta Kluge et al., 1995: 105. polyphyletic. Diagnosis. Larval maxilla with 3 (or fewer) Age: Jurassic - Recent. dentisetae (see discussion below). Euplectoptera are divided into (2.1) Pos- Probably a paraphyletic taxon, as one of teritorna and (2.2) Anteritorna. its groups (which one, is unknown) may be a sister group for Bidentiseta. 2.1. Posteritorna Kluge et al., 1995 'Tridentiseta are divided into Tetramero- tarsata Kluge, 1997 and a paraphyletic This group corresponds in its content to: group described as superfamily Siphlonuro- - Larves cryptobranches: Lestage, 19 17: 236; idea sensu Kluge et al., 1995. - Prosopistominae sensu Lameere, 1917: 74; - Prosopistomatoidea sensu Edmunds & 2.2.2. Bidentiseta Kluge, 1993 Traver, 1954b: 240; - Baetiscoidea sensu Peters & Hubbard, This group corresponds in its content to: 1989: 115; - Bidentiseta Kluge, 1993a: 41; Kluge et - Posteritorna: Kluge, 1992a: 10 (publica- al., 1995: 105. tion for temporary usage); Diagnosis. Larval maxilla with 2 (or fewer) - Posteritorna: Kluge, 1992b: 24 (publica- dentisetae (see discussion below). tion for educational usage); A holophyletic taxon. -- Posteritorna Kluge et al., 1995: 105. Bidentiseta are divided into (2.2.2.1) Bran- Diagnosis. Tornus of fore wing behind chitergaliae and (2.2.2.2) Furcatergaliae, apex of CUP(for the phylogenetic signifi- cance of 'this character see discussion below). 2.2.2.1. Branchitergaliae taxon nov. In larval maxilla, number of dentisetae in- definite, more than 3 (see discussion below). This group corresponds in its content to: In addition, a number of autapomorphies -- Setisura sensu Kluge, 1993a: 41; Kluge listed in discussion below. et al., 1995: 105 (non Setisura McCafferty, A holophyletic taxon. 1991). Posteritorna include Baetiscidae' (with a A holophyletic taxon. See apomorphies single genus Baelisca) and Prosopistomsti- discussed below. dae (with a single genus Prosopistoma). Branchitergaliae are divided into the group Eusetisura taxon nov. (which includes 2.2. Anteritorna Kluge, 1993 Isonychiidae sensu Landa, 1973, Coloburis- cidae sensu Landa, 1973 and Oligoneuriidae ?%is group corresponds in its content to: sensu Ednlunds & Traver, 1954b) and the - Larves nudibranches: Lestage, 1917: superfanlily Heptagenioidea sensu Kluge et 244; al., 1995 (which includes Pseudiron, Arthro- - Anteritorna: Kluge, 1992a: 10 (publica- plea and Heptageniidae sensu Landa, 1969a. tion for temporary usage); - Anteritorna: Kluge, 1992b: 24 (publica- 2.2.2.2. Furcatergaliae taxon nov. tion for educational usage); - Anteritorna Kluge, 1993b: 35. This group corresponds in its content to: Diagnosis. Tornus of fore wing between - Furcatergalia sensu Kluge, 1993a: 41;. apices of veins CuA and CUP(about phylo- 1993b: 41 ; Kluge et al., 19b5: 105 (non Fur- genetic significance of this character see dis- catergalia Kluge, l 989). cussion below). Larval maxilla with number A holophyletic taxon. See apomorphies of dentisetae determined, not more than 3 discussed below. (see discussion below). Furcatergaliae are divided into Ephe- A hoiophyletic taxon. meroidea sensu Edmunds & Traver, 1954b, iOOSYST. ROSSICA Vol. 7 N.Ju.Mugz: ClarsiJication of Ephemeroptera Figs ,l4.Wings of Posteritorna. 1-2, Buetisca rogersi Bern.; 3-4, Prosopistoma foliaceum Fourcr. (male). 1,3: fore wing; 2,4: its base. Caenoidea sensu Edmunds & Traver, 1954b, as PCu); CuA and CUPrun nearly parallel to Ephemerelloidea sensu Koss, 1973 and Lep- each other, without branches or intercalaries tophlebiidae. between them (Figs 1, 3). In Anteritorna, tornus is situated betueen CuA and CUP, DISCUSSION these veins are strongly divergent, with branches of CuA or intercalaries between Posteritorna and Anteritorna them (only in Ametropodidae CUPand Ar terminate verv close to the tornus'l. 'There are different views on the homology In Posteritorna, tornus (the hind angle) of of veins of Baerisca @emoulin, 1969; Ed- the fore wing is situated distinctly behind rnunds & Traver, 1954a) and Prosopisloma Cup, usually behind A 1 (which is also known (Gillies, 1954, 1956). The homologization ac- 258 N.Ju. Huge: L'lassifico~iion oJ Ephemeroptera ZOOSYST. ROSSICA Vol. 7 cepted here is based on comparison of bases Prosopistoma were regarded as not related of concave and convex veins (Figs 1-4). and were placed in different taxa. Eaton The difference in position of tornus in Pos- (1 883- 1888) placed Baetisca in his "Series I" teritorna and Anteritorna is independent of of "Group 111" (together with Siphlonurus the size of hind wings, as Anteritorna in- and others) and placed Prosopisroma in "Se- clude species with hind wings well devel- ries 111" of "Group 11" (together with Caenis oped, or rudimentary, or completely absent. and Tricorythus). Demoulin (1958, 1961) In the cases when the hind wing is rudimen- placed Baetiscidae in Oligoneurioidea and tary or absent, the tomus of fore wing is Prosopistomatidae in Ephemerelloidea; later usually shifted proximally, becomes obtuse (Demoulin, 1969), he regarded Baetiscidae or is lost, but it never changes its position to be related with Oniscigastridae, and Pro- relatively to the terminations of the longitu- sopistomatidae with Ametropodidae. Tsher- dinal veins, being in Anteritorna always be- nova (1970, 1980) placed Baetiscidae in tween CuA and CUP.The difference in posi- Ephemerelloidea, and Prosopistomatidae in tion of tornus in Posteritorna and Anteri- Caenoidea. McCafferty (1991) placed Baeti- torna can be explained by independent ori-
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