CLADOENDESIS OF EPHEMEROPTERA
Fimbriatotergaliae Fossoriae - Behningia/fg1)
Nomen hierarchicum: Behningia/fg1 [f:1937; g:1930] (incl. Protobehningia)
In circumscription fits:
— fam. Behningiidae: Tshernova & Bajkova 1960: 410
— superfam. Behningioidea: McCafferty 1991a: 349
— Behningia/fg1: Kluge 2000: 252
References. Tshernova 1970: * *; – Peters & Gillies 1991: * *; – Elpers & Tomka 1994b: *; – Kluge 1997d: *; – Kluge 2004: * * *
Autapomorphies of Behningia/fg1.
(1) Larva [burrowing – see Fossoriae (1)] has on its head, thorax and abdomen projections of unique shape which are covered by stout setae directed posteriorly, that helps to move in sand (Kluge 2004: Fig.73:A,E):
Larval frons anteriorly bears a pair of wide rounded projections which cover from above not only clypeus [see Fossoriae (2)], but also antennae bases; these projections are densely covered with stout setae directed posteriorly. Due to this, imaginal antennae are located on a pair of projections directed anteriorly (Tshernova 1938: Fig.1).
On larval pronotum ends of the crest separating collar [see Fimbriatotergaliae (1)] are transformed to a pair of large flat antero-lateral projections, densely covered with stout setae directed posteriorly.
Larval mesothoracic episterna are more or less projected anteriorly in a form of bald sclerotized projections (especially remarkable in Behningia/ /fg2), and epimera are densely covered with stout setae directed posteriorly.
Larval abdominal terga bear a pair of longitudinal ridges covered with setae; in Protobehningia these ridges are shallow (Kluge 2004: Fig.73:E), while in Behningia/fg2 they are projected laterally and cover tergalii from above (Kluge 2004: Fig.73:A-B).
In Protobehningia burrowing leg specialization is retained, but in Behningia/fg2 body shape and method of movement in sand are strongly modified, and leg specialization is changed (see below).
(2) Mouth apparatus [located at a distance from anterior margin of head because of frontal projections – see (1)] is specialized for carnivorism:
Mandible is shortened perpendicular to its axis of articulation, mola without grater, incisor and kinetodontium are stout; initial asymmetry of mandibles is either lost (in Behningia/fg3), or nearly lost (in Protobehningia and Dolania) (Elpers & Tomka 1994b:Fig.4,5,19,20).
Labium has glossae and paraglossae more or less diminished and palps enlarged, massive, without muscle inside 2nd segment (border between 2nd and 3rd segments is distinct in Dolania, and indistinct in Protobehningia and Behningia/fg3).
(3) Winged stages are short-living; moult from subimago to imago takes place in males only; middle and hind legs of male and all legs of female are non-functional (non-unique apomorphy – see Index of characters ). In female all legs are vestigial, tarsi are non-segmented, claws are lost. In male imago middle and hind legs lack tarsi; at least in Dolania (according to Peters & Peters 1986) middle and hind legs of male subimago with 2-segmented tarsus and one claw.
(4) RSa2 of fore wing is simple, not forming triad. The same in some other short-living mayflies [see (3)] (see Index of characters [2.2.37]).
(5) On fore wing bifurcation of MA is transferred to wing base. The same in some other short-living mayflies [see (3)] (see Index of characters [2.2.43]).
(6) Cubital field of fore wing [see Anteritorna (1)] with 1–8 intercalaries sometimes alternating as concave and convex, the anteriormost intercalary being the longest (non-unique apomorphy — see Index of characters [2.2.52]). CuA is either simple (in Protobehningia), or bifurcate [see Behningia/fg2 (3) below], thus no more than one vein arises from CuA.
(7) Amphitornal margin of imaginal wing bears setae similar to that of subimago. Non-unique apomorphy (see Index of characters [2.2.27]).
(8) In male imago on fore legs (elongate and specialized for grasping female, as in other mayflies) tarsus is non-segmented, with one blunt claw only (Keffermuller 1959: Pl.V; Peters & Peters 1986: Fig.1; Peters & Gillies 1991: Fig.2). Unique apomorphy.
(9) Penis lobes are very long, much longer than gonostyli [at the same time gonostyli are normally developed, not shortened – see (10)]. Unique apomorphy: besides Behningia/fg1 so long penes are present only in some Geminovenata, but there penes are protractible, while penes of Behningia/fg1 are rigid.
(10) Gonostylus [see (9)] lacks distal segments. Non-unique apomorphy (see Index of characters [2.3.12]).
(11) Imaginal and subimaginal caudalii [see (12)] lack distinct segmentation. Non-unique apomorphy, the same in some Palingenia/f3=g2 and others.
(12) In male imago paracercus [see (11)] is shortened, much shorter than cerci: 1/2 of cerci length in Protobehningia and strongly shortened in Behningia/fg2 (but in female paracercus is subequal to cerci). Non-unique apomorphy (see Index of characters [2.3.22]).
Characters of Behningia/fg1 of doubtful polarity (see Classifications of Fimbriatotergaliae II).
(13) Mandibles lack tusks. Probably result of reduction [see Fimbriatotergaliae (8)].
(14) On inner side of femur and tibia of larval hind leg spiny setae are absent. Probably result of reduction [see Fimbriatotergaliae (9)].
(15) Tergalius I [different from others – see Fimbriatotergaliae (6)] has a form of a single large (as long as tergalii of next pairs) lamella with numerous marginal processes (Kluge 2004: Fig.73:A,E). As in other Fimbriatotergaliae tergalius I is strongly diminished and always lacks processes, we can assume that in Behningia/fg1 vestige of this tergalius has been secondarily enlarged and got the same processes as on tergalii II-VII, but remains to be unilamellate.
Plesiomorphies of Behningia/fg1. Maxillary palp is always 3-segmented [in spite of specialization of mouth apparatus – see (2)] (unlike Cryptoprosternata). Furcasternal protuberances are contiguous (unlike Campsurus/fg1, Caenotergaliae and some others – see Index of characters [2.2.23]).
Size. Fore wing length 6–24 mm.
Distribution. Holarctic and Oriental Region.
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