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Ephemeridae LATREILLE, 18101 EPHEMERIDAE 437 Ephemeridae LATREILLE, 18101 In Europe represented by the subfamily Ephemerinae2 DIAGNOSIS. Rather large species of similar habitus and with the nominotypical genus Ephemera LINNAEUS colouration. Black marks on abdominal terga diagnos- only, world wide 6-8 genera including well over 50 tic, similar in larvae and imagines. Wings with black species. Three subgenera (Aethephemera, Ephe - dots, in the fore wing vein AA simple. Paracercus mera, Sinephemera)3 may be recognized. The higher present in all stages, subequal to cerci in length. classification is still in discussion (cf. McCafferty 1972, ARVAE of the burrowing type. Antennal segments 1973, 1979, 1991 and 2004, Entomol. News 115: L with whorls of fine setae. Frontal projection bifurcate. 84; Edmunds & McCafferty 1996; Kluge 2003; Klu- Mandibles with long and slender elongation (“tusk“), ge 2004, Phyl. Syst. Eph., p. 233-237) and no de- easily visible in dorsal view. Tusks slightly curved, sim- tailed cladistic analysis including larval and imaginal ple, without ramifications or secondary teeth, with characters is available at present4. The subfamily only a few inconspicuous bristles. Tibiae more or less Ephemerinae (including Ephemera LINNAEUS and broadened, claws without teeth. 7 pairs of lateral gills. Afro mera DEMOULIN) is currently defined as mono- phyletic by the following apomorphies: In larvae Gill 1 small, bifurcate tubiform, the others deeply cleft mandibular tusks are curved, without denticles and with fringed margins. round in cross section. Abdominal segments 7-9 elon- IMAGINES of characteristic habitus. In male imagines gated posteriorly, gills inserted in the middle of the both tarsal claws blunt (one blunt, one hooked in segments. Gill 1 vestigial. In imagines fore wing with subimagines). In hind wings vein MP2 usually attached several veins from AA to hind margin of wing. Man - basally to vein CuA. Black marks on abdominal terga dibular tusks and their function have been studied in usually diagnostic. Ground pattern of black dots on detail by Keltner & McCafferty (1986), Bae & McCaf- wings may be used as an additional character for iden- ferty (1995) and Edmunds & McCafferty (1996). tification but shows considerable intraspecific varia- Distribution of Ephemeridae is worldwide except tion. Male genitalia rather similar throughout the Austra lia, some biogeographical aspects have been subgenus. Penes simple, with slender spine-like me- discussed by McCafferty (1973, Oriental Insects 7: dian titillators (missing in subgenus Sinephemera). 49, McCafferty (1999) and Edmunds & McCafferty Forceps four-segmented, with apical and subapical (1996). Species diversity of Ephemerinae is highest segments short. Eggs with rather smooth extrachori- in the East Palaearctic and Oriental Region (and tran- onic adhesive layer covering the chorionic sculpturing sitory areas). For diagnosis and biology see Ephe - (see Ubero-Pascal & Puig 2007, Zootaxa 1465: 15– mera. 29, fig. 3). DISTRIBUTION. Holarctic, Oriental and Afrotropic re- Ephemera LINNAEUS, 1758 gion5. Ephemera LINNAEUS, 1758; Syst. Nat., ed. 12: 546 BIOLOGY. Larvae may occur in almost all types of freshwater with sandy or silty bottom. Some species Type species: Ephemera vulgata LINNAEUS, 1758. (e.g. Ephemera vulgata and E. glaucops) inhabit still- SubseQuent designation by Westwood (1836, Parting- waters including backwaters, lakes and oligotrophic ton’s British Cyclopaedia 2, p. 439). fish-ponds. They live in imperfectly U-shaped tubes Nirvius NAVÁS, 1922; Bol. Soc. Entomol. Espagna 5: 56 in the substrate and feed on detritus, filtering the [junior subjective synonym teste Lestage (1922, Bull. Soc. water current generated by synchronized gill move- Entomol. France 16: 253)] ment (active filter feeding). Larval populations (includ- 1 For authorship of the family see Peters & Hubbard (1989, J. New York Entomol. Soc. 97,1: 115). 2 A second (monotypic) subfamily Ichthybotinae KOSS & EDMUNDS, 1974 (Zool. J. Linn. Soc. 55: 295) is restricted to New Zealand (s.a. Demoulin 1957, Bull. Ann. Soc. Roy. Entomol. Belg. 93: 335). 3 Aethephemera MCCAFFERTY & EDMUNDS, 1973; Pan-Pac. Entomol. 49 (4): 306 (India). Dicrephemera MCCAFFERTY & EDMUNDS, 1973; Pan-Pac. Entomol. 49, 4: 302 (India), originally proposed as a subgenus of Ephemera LINNAEUS, is currently considered a junior subjective synonym of Afromera DEMOULIN, 1955 (teste McCafferty & Edmunds 1979, Aquatic Insects 1, 3: 170). 4 McCafferty (1972, J. Georgia Entomol. Soc. 7: 51) placed the Nearctic genus Pentagenia WALSH (1863, Proc. Entomol. Soc. Philadelphia 2: 196) in a separate family Pentageniidae. Taxa of Pentageniidae have subsequently been transferred to Ephemeridae (Edmunds et al. 1976, Mayflies North Central Amer., p. 281, see also p. 292 footnote) and later to Palingeniidae (Edmunds & Mc- Cafferty 1996: 73, see also p. 446). McCafferty (1991, Ann. Entomol Soc. Amer. 84: 343) established the subfamily Hexageniinae to accommodate the related Nearctic and Neotropic genera Hexagenia WALSH and Litobrancha MCCAFFERTY together with the Afro-Oriental genera Eatonigenia ULMER and Eatonica NAVÁS. 5 At present Ephemera mooiana MCCAFFERTY, 1971 (J. Entomol. Soc. South Afr. 34: 57-62) is the only known representative in the Afrotropics. 438 EPHEMERIDAE ing Nearctic and Oriental species) are usually invaded presence of a concave frontal process, vestigial man - by numerous larvae of the chironomid Epoicocladius dibular tusks subeQual to incisors (Barber-James & ephemerae. This phenomenon is usually considered Lugo-Ortiz 2003: 80, fig. 2.73) and a Quite uniQue a mere commensalism but the relationship to the host mesonotal structure (for details see Kluge 2004: 236 is probably more complicated (see Tokeshi 1986 and and fig. 71 F). Imagines differ in smaller hind wings Svensson 1980 for details). Developmental cycle (de- (with basitornal margin shorter) and different shape pendent on water temperature) lasts from 1 to 3 of titillators (missing in some species). years. Under suitable conditions (e.g. chalkstreams, lake shores) high population densities are observed EXTRALIMITAL SPECIES: and characteristic mass emergence takes place, when Ephemera brunnea HUBBARD & PETERS, 1978; Oriental In- imagines are found only during a short time of the sects Suppl. 9: 15 [new replacement name for E. striata year. Under less favourable conditions small numbers ALI = Ephemera striatum [sic] ALI, 1970 (Pakistan J. Sci. of fully grown nymphs may be found throughout the 22:120) nec Ephemera striata LINNÈ, 1767] nom. pre- summer months (May-October). Subimagines emerge occ. (Pakistan [? subgeneric placement]) on the water surface. Males swarm near the shore and Ephemera formosana ULMER, 1920; Arch. Naturg. 85A oviposition takes place on the water surface. (11): 6 (Taiwan [? subgeneric placement]) Ephemera (Ephemera) hainanensis YOU & GUI, 1995; REMARKS. Identification rests mostly on abdominal Econ. Insect Fauna China 48: 99 (China) markings and male genitalia. Both of them are some- Ephemera (Ephemera) hongjiangensis YOU & GUI, 1995; times difficult to interpret and show considerable in- Econ. Insect Fauna China 48: 98 (China) traspecific variation. Bleached specimens (especially Ephemera (Ephemera) hunanensis YOU & GUI, 1995; Econ. Insect Fauna China 48: 104 (China) females) and cast exuviae usually not identifiable on Ephemera hsui YOU & GUI, 1995; Econ. Insect Fauna the species level. In freshly moulted specimens the China 48: 104 (China [? subgeneric placement]) colouration pattern is but weakly developed and may Ephemera (Ephemera) iwatensis MATSUMURA, 1931; 6,000 sometimes be misleading. The same applies to heavily Ill. Ins. Jap. Empire, p.: 1467 (Japan) parasitized specimens ( Thomas & Parpet 2005, Ephemera (Sinephemera) japonica MCLACHLAN, 1875; Ephemera 6, 1: 8). No recent revision for the genus Trans. Entomol. Soc. London 1875: 169 (Japan) is available, a world check list has recently been pro- Ephemera (Ephemera) jianfengensis YOU & GUI, 1995; vided by Hwang et al. (2008). Palaearctic taxa (imag- Econ. Insect Fauna China 48: 102 (China) ines and larvae [so far known]) have been revised and Ephemera (Ephemera) kirinensis HSU, 1931; Peking Nat. Hist. Bull. 6 (2): 40 (China) keyed by Tshernova (1973, Entomol. Obozr. 52, 2: Ephemera (Sinephemera) nigroptera ZHOU, GUI & SU, 223-233). Most European taxa have been keyed by 1998; Entomol. sinica 5 (2): 139 (Ahui Prov. China) Jacob et al. (1975, Entomol. Nachr. Ber. Dresden 19: comb. nov. [new subgeneric combination] 186) and Bauernfeind & Humpesch (2001). Eastern Ephemera (Ephemera) orientalis MCLACHLAN, 1875; Palaearctic taxa have been keyed by Tshernova et al. Trans. Entomol. Soc. London 1875: 167 (Japan (1986, pp. 105-106) and Kluge (1997), for Japanese [=Ephemera amurensis NAVÁS, 1912; Rev. Russ. Ento- and Korean taxa see Kuroda & Watanabe (1984, Ka- mol. 12: 414 (Siberia) = Ephemera modesta BRODSKY, gawa Seibutsu 12: 24), Yoon & Bae (1985, Entomol. 1930; Rossiyskoe Entomol. Obozr. 24 (1-2): 31 teste Res. Bull. 11: 93-109), Ishiwata (2003), Bae (1997) Tshernova (1952, Trudy Amur. Ikht. Eksped. 1945-49, and Bae et al. (1994). Nearctic species have been 3: 234)] = Ephemera kuccharonis MATSUMURA, 1931, 6,000 Ill. Ins. Jap. Empire, p. 1468 (Japan) keyed by McCafferty (1975, Trans Amer. Entomol. Ephemera pictipennis ULMER, 1924; Konowia 3: 28 (China Soc. 101: 447-504), Afrotropic taxa by McCafferty [? subgeneric placement]) (1971, J. Entomol Soc. South Africa 34: 57-62), De- Ephemera pictiventris MCLACHLAN, 1894; Ann. Mag.
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