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Keys to the known Recent species of the mayfly family Behningiidae are presented. Also included is a catalog of references to the genera and species of the Behningiidae, along with indications of described stages and geographical distributions.
The mayfly family Behningiidae is one of the smallest families of the Ephemeroptera. It comprises only four genera: the three Recent genera Behningia Motas & Bacesco, Dolania Edmunds & Traver, and Protobehningia Tshernova, and the Jurassic fossil genus Archaeobehningia Tshernova. To date there are only seven described species in the family.
Although McCafferty (1991) recently elevated the Behningiidae to its own superfamily, he presented little data (McCafferty 1979, McCafferty & Edmunds 1976) to support his rationale and his arguments therefore remain unconvincing. I still consider the family to belong in the Ephemeroidea, and accordingly, following standard practice among mayfly workers, I have left it placed therein.
Keys to the known stages of the nymphs and imagos of the Recent species of the Behningiidae are presented.
The catalog is intended to be a comprehensive, but certainly not exhaustive, listing of references in the scientific literature to all known species of the Behningiidae. Included with each species are indications of the geographic distributions listed in the scientific literature. Listed with the references to taxonomic papers are indications of the stage described or figured. Only selected taxonomic references are given for each genus.
This paper is one in a continuing series of catalogs of the Ephemeroptera (c.f. Hubbard & Pescador 1978, Hubbard & Peters 1978, Hubbard 1979, 1982a, 1982b, 1986, 1987, 1990, Hubbard & Savage 1981, Hubbard, Dominguez & Pescador 1992).
Archaeobehningia Tshernova, 1977, 1977:94; Soldán, 1979, 72:636; Kaludina, 1980, 178:229; Tshernova, 1980, 178:33; Landa & Soldán, 1985, 4:99; Hubbard, 1987, 129:61; Peters & Gillies, 1991, p:211.
Type-species: Archaeobehningia edmundsi Tshernova, original designation.
Archaeobehningia edmundsi Tshernova, 1977, 1977:95, fig. 2, pl. 10 fig. 2 (nymph); Sinitshenkova, 1985, 211:22; Hubbard, 1987, 129:62; Peters & Gillies, 1991, p:211.
Archaebehningia [sic] edmundsi; Sinitshenkova, 1984, p:63.
Distribution: Jurassic of Russian Transbaikal (fossil).
Behningia Lestage, 1930, 69:436; Tshernova, 1938, 1938:131; Lestage, 1938, 78:315; Tshernova, 1940,1:130; Bertrand, 1954, p:254; Edmunds & Traver, 1959, 52:46; Tshernova & Bajkova, 1960, 39:415; Landa, 1969, 18:295; McCafferty, 1975, 101:451; Soldán, 1979, 72:636; Landa & Soldán, 1985, 4:99; Peters & Peters, 1986, 69:246; Tshernova et al. 1986, 1:111; Peters & Gillies, 1991, p:211.
Type-species: Behningia ulmeri Lestage, monotypy.
Behningia lestagei Motas & Bacesco, 1937, 24:27, fig. 1-2 (nymph); Lestage, 1938, 78:315 (nymph); Motas & Bacesco, 1940, 26:78, figs. 1-4, pl. 1 (nymph); Demoulin, 1952, 28(21):1; Demoulin, 1955, 91:207; Keffermüller, 1957, 7:254; Edmunds & Traver, 1959, 52:46, fig. 13-23 (nymph); Poprawska, 1960, 4:160; Tshernova & Bajkova, 1960, 39:410; Keffermüller, 1963, 12:323; Brodsky, 1974, 53:296, fig. 1g (adult); Koss & Edmunds, 1974, 55:324, pl. 12 fig. 138-139 (egg); Peters & Peters, 1977, 62:409; Tshernova, 1977, 1977:95; Soldán, 1979, 72:636; Puthz, 1978, 2:263; Fink, Soldán, Peters & Peters, 1991, 69:1083.
[?] Behningia lestagei; Bertrand, 1954, p:254, fig. 123 (nymph).
Behningia lestagei ?; Keffermüller, 1959, 19(5):8, pl. 4-7, 9-12 (male, female, nymph); Keffermüller, 1960, 19(8):6; Peters & Peters, 1986, 69:245.
Behningia ulmeri; Landa, 1969, 18:295, pl. 28 (male, female, nymph).
Behningia ulmeri [partim]; Kluge, 1989, pl. 5 fig.3-4 (nymph).
Distribution: Moldavia, Poland, Czechoslovakia, Romania.
Behningia ulmeri (?); Tshernova, 1938, 1938:132, fig. 1-4 (male).
Behningia ulmeri [partim]; Tshernova, 1940, 1:133; Kluge, 1989, pl. 5 fig.3-4 (nymph).
Behningia ulmeri ?; Demoulin, 1952, 28(21):1, fig. 5 (male); Tshernova, 1952, 3:248, figs. 20-24 (nymph); Tshernova, 1958, 37:73.
Behningia ulmeri; Bertrand, 1954, p:254, fig. 124 (male).
Behningia tshernovae Edmunds & Traver, 1959, 52:47, fig. 23-30 (male, nymph); Landa, 1969, 18:297; Peters & Peters, 1977, 62:409; Tshernova, 1977, 1977:95; Tiunova, 1986, p:21; Tshernova et al. 1986, 1:111, fig. 48(1-4) (male).
"Nov. gen., nov. sp. ?" Behning, 1924, 1:252, fig. 33-34 (nymph).
"Merkwürdige Ephemeriden-Nymphe" Ulmer, 1924, 7:3, fig. 1-6 (nymph).
Behningia ulmeri Lestage, 1930, 69:436; Motas & Bacesco, 1937, 24:25 (nymph); Lestage, 1938, 78:315 (nymph); Motas & Bacesco, 1940, 26:78 (nymph); Demoulin, 1952, 28(21):1; Demoulin, 1955, 91:207; Keffermüller, 1957, 7:254; Edmunds & Traver, 1959, 52:46, fig. 22; Tshernova & Bajkova, 1960, 39:410; Peters & Peters, 1977, 62:410; Tshernova, 1977, 1977:95; Soldán, 1979, 72:636; Puthz, 1978, 2:263; Peters & Gillies, 1991, fig. 9, 12-13 (male).
Behningia ulmeri ?; Klyutschareva, 1963, Zoologicheskii Zhurnal 42:1604.
[not] Behningia ulmeri; Tshernova, 1938, 1938:132, fig. 1-4; Bertrand, 1954, p:254, fig. 124 (male); Bertrand, 1954, p:254; Landa, 1969, 18:295, pl. 28 (male, female, nymph).
[not] Behningia ulmeri ?; Demoulin, 1952, 28(21):1, fig. 5 (male); Tshernova, 1952, 3:248, figs. 20-24 (nymph); Tshernova, 1958, 37:73.
Dolania Edmunds & Traver, 1959, 52:46; Tshernova & Bajkova, 1960, 39:415; McCafferty, 1975, 101:451; Edmunds, Jensen & Berner, 1976, p:274; Berner, 1977, 22(1):48; Peters & Peters, 1977, 62:409; Landa & Soldán, 1985, S4:99; Berner & Pescador, 1988, p:259; Peters & Gillies, 1991, p:211.
Type-species: Dolania americana Edmunds & Traver, original designation.
Dolania americana Edmunds & Traver, 1959, 52:46, figs. 1-16, 31-32 (nymph); Tshernova & Bajkova, 1960, 39:414; Edmunds, 1962, 12(5):17; Schneider, 1966, 29:205; Peters & Jones, 1973, p:246; Koss & Edmunds, 1974, 55, pl. 12 fig. 140-141 (egg); McCafferty, 1975, 101:451, fig. 1 (nymph); Edmunds, Jensen & Berner, 1976, p:276, fig. 42, 197, 202, 204, 424 (male, female, nymph); Berner, 1977, 22(1):48; Peters & Peters, 1977, 62:409; Tshernova, 1977, 1977:95; Soldán, 1979, 72:636, fig. 1-15 (male, female, nymph); Tsui & Hubbard, 1979, 67:119; Finn & Herlong, 1980, 91:102; Harvey, Vannote & Sweeney, 1980, p:211; Dakin & Felder, 1981, 64:454; Peters, 1982, 6:28; Sweeney & Vannote, 1982, 36:811; Basha & Pescador, 1984, p:205; Benke et al., 1984, 54:43; Fink, 1986, p:1; Peters & Peters, 1986, 69:245, fig. 1-3 (male, female); Peters, Peters & Fink, 1987, 65:3177; Berner & Pescador, 1988, p:259, pl. 14, fig. 14 (male, female, nymph, egg); Fink & Yasui, 1988, 17:448, fig. 1-9 (sperm); Peters & Peters, 1988, 97(5):8, 3 figs (male); Kluge, 1989, pl. 5 fig. 2 (nymph); Jacobs, 1990, 101:219; Fink, Soldán, Peters & Peters, 1991, 69:1083, fig. 3a-g, 6a-j, 7a-d (nymph); Peters & Gillies, 1991, p: fig. 8, 10-11 (male); Sweeney & Funk, 1991, 13:17; Peters & Peters, (in press).
Distribution: United States (Florida, Georgia, Lousiana, North Carolina, South Carolina, Wisconsin-Minnesota).
Dolania sp. nov.; Sweeney & Funk, 1991, 13:18.
Protobehningia Tshernova in Tshernova & Bajkova, 1960, 39:411; Peters & Peters, 1977, 62:410; Soldán, 1979, 72:636; Landa & Soldán, 1985, 4:99; Tshernova et al., 1986, 1:111; Peters & Peters, 1986, 69:246; Peters & Gillies, 1991, p:207.
Type-species: Protobehningia asiatica Tshernova, original designation.
Protobehningia asiatica Tshernova in Tshernova & Bajkova, 1960, 39:413, fig. 1-3 (nymph); Peters & Peters, 1977, 62:410; Tshernova, 1977, 1977:95; Tiunova, 1986, p:21; Tshernova et al., 1986, 1:111; Kluge, 1989, pl. 5 fig. 1 (nymph); Peters & Gillies, 1991, p:207, fig. 7 (nymph).
Protobehningia merga Peters & Gillies, 1991, p:208, fig. A, 1-6 (male, nymphal exuviae).
I thank Janice G. Peters, Department of Entomology, Florida A&M University, for her helpful comments.
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_____. (in press). Evidence for a dispersal morph in females of Dolania americana (Ephemeroptera: Behningiidae). Proceedings of the VIIth International Conference on Ephemeroptera, Sandhill Crane Press, Gainesville, Florida.
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