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Mayflies of Uncertain Systematic Position Appendix Mayflies of uncertain systematic position Most fossil (mainly Mesozoic and Palaeogenous) Anteritoma INCERTAE SEDIS 1 and some insufficiently described Recent mayflies t Hexagenites/fgl have an uncertain position among Ephemeroptera. Nomen hierarchicum: Hexagenites/fgl [f: Hexagenitinae Here generic and species names belonging to such Lameere 1917: 74; g: Hexagenites Scudder 1880: 6, taxa are accounted in the following method: typus H weyenberghii Scudder 1880 (design. orig.)J; Anteritoma INCERT AE SEDIS - names given for synn. subj.: Paedephemeralfg [f: Paedephemeridae fossil supraspecies taxa and selected Recent species Lameere 1917: 49; g: Paedephemera Handlirsch 1906: 601, typus Ephemera multinervosa Oppenheim 1888 of mayflies, whose anteritomous wing structure is (design. Tshemova 1961: 859)]; Stenodicranumlfg [f: known for certain. Stenodicranidae Demoulin 1954f: 553; g: Stenodicra­ Euplectoptera INCERTAE SEDIS (p.356) - names num Demoulin 1954f: 571, typus Ephemera cellulosa given for fossil mayflies known as larvae or adults Hagen 1862 (design. orig.)] [synn. subj.: cellulosa [£.] with poorly preserved wings. Probably, most or all = weyenberghii [H] = multinervosa [£.] (Demoulin 1970c: 7)]. of these mayflies belong to Anteritoma, but this is In circumscription matches: not proven because wing venation and number of - subfam. Ephemeropsinae Cockerell 1924: 136; dentisetae are unknown. - fam. Hexagenitidae: Tshemova 1961: 589. Euephemeroptera INCERTAE SEDIS (p.359) - Mesozoic taxa, one of which (Litophlebialfgl) has References. Cockerell 1924: tl $; - Tshemova 1961: $*; Meshkova 1961: tl; - Tshemova 1970: tl $ - unusual wing structure and others which were Tshemova& Sinitshenkova 1974: $'; - McCafferty 1990: established for indeterminable fragments. tl'; - Martins-Neto 1996: tl'. Pterygota INCERTAE SEDIS (p.360) - Palaeozoic Autapomorphies of Hexagenites/fgl. taxon Syntonopteralfgl possibly related to Epheme­ (I) On fore wing CuA bifurcates to CuAl and roptera s.l., and some taxa erroneously attributed to "CuA2" (non-unique character - see Index of Ephemeroptera. characters [2.2.50]), from the bifurcation a vein Euarthropoda and Animalia INCERT AE SEDIS "iCu" arises, which forms several (3-5) triads (p.361) - fossils, which some authors regard to following one another: anterior branch of each triad belong to mayfly larvae. forms next triad; each of these triads has anterior Nomina dubia (p.361) - old forgotten species branch arched by its convexity anteriorly; all names, possibly belonging to well-known species. branches of these triads go to basitomal margin of wing. Unique apomorphy. (2) Fore and hind wings with more or less expressed vein gemination: at least there are pairs of veins brought together: RSp+MAh iMA+MA2, MP2+CuA; MPI and iMP are more or less approxi­ mated to iMA+MA2• Non-unique apomorphy (see Index of characters [2.2.28]). (3) Tergalius VII is enlarged, distinctly longer than others, while tergalii I-VI have subequallength (unique apomorphy). On all tergalii at least costal rib is thickened and anal rib is situated at a distance 353 354 ApPENDIX from anal margin (as in some other mayflies - see [E.]: Ea; selini [H.]: Ea; trisetalis [E.]: 19, Ea. Index of characters [1.3.28]). Plesiomorpbies of Hexageniteslfgl. Larva has *** siphlonuroid appearance, with dense primary swimming setae on caudalii. Other Anteritoma INCERTAE SEDIS Size. Fore wing length 16-42 IDID. Age. Late Jurassic - Late Cretaceous (Europe, Anteritorna INCERTAE SEDIS 2. t Cronicus/g [g: Croni­ Asia, South America). cus Eaton 1871:133, typus Baetis anomala Pictet & Hagen 1856 (monotypy; syn.subj.: Baetis longipes Pictet & Hagen Nominal taxa included. Hexageniteslfgl includes: 1856)]. (1) Ephemeropsis/fg [f: Ephemeropsinae Cockerell 1924: Reference. Demoulin 1968a: 0 Ea. 136; g: Ephemeropsis Eichwald 1864: 21, typus E. Characters of unclear phylogenetic status. trisetalis Eichwald 1864 (monotypy)]; (2) Phacelobran­ (1) On hind wing bifurcation of MP2 is transferred chus/g [g: Phacelobranchus Handlirsch 1906: 604, typus distally or lost (see Index of characters [2.2.69]). Ph. braueri Handlirsch 1906 (monotypy)], regarded as a (2) Imaginal paracercus is vestigial (see Index of generic synonym of Ephemeropsis (Tshemova 1961: 861); characters [2.3.20]). (3) Protoligoneurialg [g: Protoligoneuria Demoulin (3) Gonostylus with 3 distal segments (see Index of 1955e: 271, typus P. Iimai Demoulin 1955 (design. orig.); characters [2.3.12]). (4) Hexameropsis/g [g: Hexameropsis Tshernova& Sini­ Plesiomorphies. Fore wing with several (5-6) tshenkova 1974: 132, typus H. selini Tshemova & Sini­ simple or branched veins going from CuA to basitomal tshenkova 1974 (design. orig.)]; (5) Mongologenites/g [g: and tomoapical margins. Imaginal and subimaginal claws Mongologenites Sinitshenkova 1986: 45, typus M are ephemeropteroid. lagueatus Sinitshenkova 1986 (design. orig.)]; (6) Palaeo­ Age. Eocene (Baltic amber). baetodes/g [g: Palaeobaetodes Brito 1987: 594, typus P. Material examined: -. costalimai Brito 1987]; (7) Cratogenites/g [g: Cratogeni­ Comment. Demoulin (1968a) attributed Cronicus to tes Martins-Neto 1996: 180, typus C. corratiniae Martins­ Coloburiscinae (i. e. ColoburiscuS/fgl) basing on plesio­ Neto 1996 (design.orig.)]; (8) Cratogenitoides/g [g: morphies only. In order to clarify its systematic position, Cratogenitoides Martins-Neto 1996: 181, typus C. delclosi thorax structure of types (deposited in Humboldt Univer­ Martins-Neto 1996 (design. orig.)]; (9) Caenoephe­ sity) should be examined. meralg [g: Caenoephemera Lin & Huang 2001: 748, typus C. shanguanensis Lin & Huang 2001 (design.orig.)]. Anteritorna INCERTAE SEDIS 3. t Siphlurites/g [g: Siph­ Possibly, here also belongs Costalimella (see below, lurites Cockerell 1923: 170, typus S. explanatus Cockerell Euplectoptera INCERTAE SEDIS). 1923 (design. orig.)]. Species composition of Hexagenites/fgl. Following Reference. Demoulin 1970d: fore wing. formally described species belong here for certain: From Character of unclear phylogenetic status. Mesozoic of Europe: cellulosa Hagen 1862 [Ephemera] (1) Cubital field offore wing with 2 veins going from (synn. subj.: mortua Hagen 1862 [Ephemera], weyen­ CuA to basitomal margin, 1 intercalary going to berghii Scudder 1880 [Hexagenites], multinervosa tomoapical margin, and several (9-10) simple or branched Oppenheim 1888 [Ephemera], oppenheimi Handlirsch veins going from this intercalary to basitornal margin (see 1906 [PaedephemeraD. From Mesozoic of Siberia, Index of characters [2.2.51]). In other respects wing Mongolia and China: lagueatus Sinitshenkova 1986 venation is plesiomorphic. [Mongologenites], martynovae Tshemova 1961 [Epheme­ Age. Miocene (USA). ropsis], selini Tshemova & Sinitshenkova 1974 [Hexa­ Material examined: -. meropsis], shanguanensis Lin & Huang 2001 [Caeno­ ephemera], trisetalis Eichwald 1864 [Ephemeropsis] Anteritorna INCERTAE SEDIS 4. t Electrogenialg [g: (synn. subj.: rossica Eichwald 1866 [Lethaea] , orientalis Electrogenia Demoulin 1956a: 95, typus E. dewalschei Eichwald 1868 [Estheria], braueri Handlirsch 1906 Demoulin 1956 (design. orig.)]. [Phacelobranchus], middendorji Handlirsch 1906 [Ephe­ Reference. Demoulin 1956a: @. meropsis], melanurus Cockerell 1924 [Ephemeropsis], Characters of unclear phylogenetic status. reisi Handlirsch 1939 [Ephemeropsis]). From Lower (1) Cubital field of fore wing with 2 pairs (4 ones) Cretaceous of Brazil: britoi Martins-Neto 1996 [Palaeo­ intercalaries, among which hind pair is longer. The same baetodes], costalimai Brito 1987 [Palaeobaetodes], in Heptagennota and some others (see Index of characters corratiniae Martins-Neto 1996 [Cratogenites], delclosi [2.2.51-52]). Martins-Neto 1996 [Cratogenitoides], limai Demoulin (2) Imaginal paracercus is vestigial (see Index of 1955 [Protoligoneuria]. characters [2.3.20]). Material examined (Paleontol. Inst.): limai [P.]: 19; Plesiomorphies. 1st tarsal segment is fused with lagueatus [M.]: Ea, 19 (presumably associated); martynovae tibia (in contrast to Heptagennota-Pentamerotarsata). MAYFLIES OF UNCERTAIN SYSTEMATIC POSITION 355 Imaginal claws of all legs are ephemeropteroid. Gono­ systematic position, thorax structure of the holotype stylus with 2 distal segments. (deposited in Staatl. Mus. Naturk. Stuttgart) should be Age. Eocene (Baltic amber). examined. Material examined:-. Comment. Demoulin (1956a) attributed Electrogenia Anteritorna INCERTAE SEDIS 8. t Xenophlebia/g [g: Xe­ to Arthropleinae (i. e. Arthroplealfgl); this is wrong, as nophlebia Demoulin 1968a: 267, typus X aenigmatica Electrogenia has no articulation of 1SI tarsal segment with Demoulin 1967 (design. orig.)]. tibia [see Pentamerotarsata (1)]. In order to clarify its Reference. Demoulin 1968a: EfJ. systematic position, thorax structure should be examined. Character of unclear phylogenetic status. (1) Hind wing is diminished; cubital field offore wing Anteritorna INCERTAE SEDIS 5. t Stackelbergisca/g [g: is short, with 2 intercalaries (see Index of characters Stackelbergisca Tshernova 1967: 323, typus S. sibirica [2.2.51-52]). Tshernova 1967 (design. orig.)]. Plesiomorphies. Imaginal claws are ephemeropte- Reference. Tshernova 1967: fore wing, (9 (presu­ roid. Gonostylus with 2 distal segments. mablyassociated). Age. Eocene (Baltic amber). Plesiomorphy. Cubital field offore wing with many Material examined:-. (about 15) veins going from CuA to basitornal margin. Comment. Demoulin (l968a) attributed Xenophlebia Age. Jurassic (Siberia). to Leptophlebiidae (i. e. Leptophlebialfgl).
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