CLADOENDESIS OF EPHEMEROPTERA

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Autapomorphies of Hermanellandria.

(1) Each of two penis apices is produced into a slender sclerotized tubular process – telopenis, inside which the genital duct passes; apex of telopenis is obliquely truncated and pointed. In imago telopenes are sharply bent at their bases and/or shifted from apices of penis lobes, so that are directed either medially (in most Thraulodes), or laterally (in Farrodes), or ventrally (in Hermanellognatha), or even ventrtally-anteriorly, i.e. toward penis base. Unlike imago, in mature larva telopenes represent apical continuations of penis lobes, being directed caudally, so that gonopores occupy apical position; larval protopenis has developed gonopores with ducts lined by larval cuticle (Kluge 2008: Fig.9–11, 15, 27–28, 32, 34; Kluge 2020: Figs 106–109, 347, 500–501). In subimago telopenes either have the same position as in imago, or position intermediate between larval and imaginal ones (Kluge 2008: Fig.12). Subimaginal telopenes are sclerotized, while the rest penis is membranous.

In Hermanellognatha and Simothraulopsis each telopenis represents a tube with apical opening. In all Thraulodes each telopenis is a roll, i.e. a tube cleft up to the base. In Farrodes telopenes lack gonoducts, and gonopores are locate at their bases; probably, this is a secondary condition.

Telopenes can be confused with superficially similar processes, which occur in some other taxa and arise from penis apex toward its base (see Index of characters [2.3.16]); but unlike telopenes, penial processes of other Leptophlebia/fg1 do not bear genital ducts and in larval stage develop in the same position as in imago.

Characters of Hermanellandria of unclear phylogenetic status. 

Plesiomorphies of Hermanellandria. Glossae are small, not curved ventrally (see Index of characters [1.1.50]). On fore wing fork of MA is asymmetrical [see Leptophlebia/fg1 (5)] (see Index of characters [2.2.43]). In cubital field of fore wing anteriormost of two intercalaries [see Leptophlebia/fg1 (7)] is either basally connected with CuA, or free, but never connected with CuP (see Index of characters [2.2.51]).