CLADOENDESIS OF EPHEMEROPTERA

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Autapomorphies of Crassabwa/g1.

(1) Labial palp with 2nd segment forming prominent inner-apical projection; 2nd segment retains muscle-adductor of 3rd segment (Kluge et al. 2017: Fig.5; Kluge et al. 2018: Fig.5, 69). Similar projection independently appeared in some other taxa: among Protopatellata, in Bugilliesia (belonging to Rhithrocloeoninae) and Indocloeon; among Anteropatellata, in Cheleocloeon and some of Baetovectata (see Index of characters [1.1.58]).

(2) Larval femur on outer side with one row of stout elongate spatulate colorless setae; near apex of femur one subapical seta is locates, which has the same structure as other setae of this row, but usually is somewhat shorter and separated from nearest seta by a distance greater than other distances between setae of this row (Kluge et al. 2017: Fig.10–13, 64, 67, 125; Kluge et al. 2018: Fig.18, 91-93). In most other Baetidae larval femur either has two subapical setae brought together, or no recognizable subapical seta.

(3) Larval tibia without outer-apical seta (Kluge et al. 2017: Fig.10–12, 64, 67, 122, 125); in contrast to many other Baetidae, whose larval tibia bears one small stout seta on outer side near joining with tarsus (see Index of characters [1.2.17]). Outer side of tibia and tarsus have no other stout setae; inner side of tibia and tarsus with sparse spine-like setae (as in other taxa).

(4) Larval claw with two regular rows of denticles, among which two most distal denticles of each row are enlarged and shifted toward sides of claw. These two rows of denticles (anterior and posterior rows) are located symmetrically on inner side of the claw; usually they are also symmetric regarding size of their denticles (Kluge et al. 2017: Fig.66, 95, 126; Kluge et al. 2018: Fig.19, 70), but in flavum [Centroptilum] two distal denticles of the anterior row are much larger than two distal denticles of the posterior row (Kluge et al. 2017: Fig.15, 18–19). On individual claws instead of two large distal denticles, there is larger number of thinner denticles. Besides Crassabwa/g1, two enlarged distal denticles in each of two rows are found in some other taxa, e.g. in selected species of Cheleocloeon (Kluge 2016: Fig.23,27) and in undescribed species of Indocloeon.

(5) Egg chorion with net-like relief, whose longitudinal ridges are more wide and convex than transverse ridges, so that at least in middle part of egg this relief looks as longitudinal striation (Kluge et al. 2017: Fig.45–50, 108–113; Kluge et al. 2018: Fig.58-61,  ##; ). In many other Baetidae chorion has even net-like relief with isodiametric cells separated by equally developed longitudinal and transverse ridges.

Plesiomorphies and characters of Crassabwa/g1 of unclear phylogenetic status. 

(6) Bases of antennae are widely separated, frons between them is flat (Kluge & al. 2017: Fig.3; Kluge et al. 2018: Fig.65) (unlike carinate in some other taxa).

(7) Larval frontal suture elongate (Kluge et al. 2017: Fig. 61; Kluge et al. 2018: Fig.65).

(8) Labrum nearly semicircular with median incision; dorsal surface with pair of submedian setae and one or two pairs of latero-distal setae (Kluge et al. 2017: Figs 71, 129; Kluge et al. 2018: Fig.6, 65).

(9) Mandibles with incisor and kinetodontium fused at most length; left prostheca with 4 blunt processes and 3–5 small pointed processes more or less fused together; right prostheca nearly parallel-sided, apically divided to several processes; setae between prostheca and mola present at least on right mandible, in ludmilae [Crassabwa] on both mandibles; on left mandible molar processes much shorter than distal molar projection (Kluge et al. 2017: Figs 8–9, 69–70, 76–82, 127–128; Kluge 2018: Fig.8-9, 67-68).

(10) Maxilla has all 3 canines and 3 dentisetae; distal dentiseta much thicker than others, but bent at the same direction as others; among several setae proximad of dentisetae, 1st one either simple as others, or bifid, resembling dentiseta and possibly consists of two setae fused at base (Kluge et al. 2017: Fig.6–7; Kluge et al. 2018: Fig.7, 66).

(11) Maxillary palp 2-segmented; in Crassabwa/g2 2nd segment thick, with short and narrow apical projection, which possibly represents vestige of 3rd segment (Kluge et al. 2017: Fig.72, 130); in Susua with 2nd segment is very soft and delicate.

(12) Labium with paraglossa slightly wider than glossa (Kluge et al. 2017: Fig.5; Kluge et al.2018: Fig.5, 69). Glossa with following setal rows: median row (regular row of spine-like setae along median margin); dorso-lateral row (regular row along lateral margin, with setal bases visible in dorsal view); ventral side of glossa bears setae directed ventrally, which are located irregularly in the proximal part of the glossa and form a ventro-median row (sparse row parallel to median margin). Paraglossa with following setal rows: latero-apical setae (longest ones) form a regular sparse row along lateral margin, more densely and irregularly located on apical margin, and irregularly located on dorsal side; ventro-median row (regular row of long slender straight setae attached on ventral side and directed mainly medially); dorso-median row (regular row of thicker seta attached on dorsal side and directed medio-distally, limited by distal half of paraglossa). The same in many other taxa.

(13) Patella-tibial suture typical for Protopatellata: in larvae of both sexes and in female at all stages (larva, subimago and imago) fore tibia without patella-tibial suture, middle and hind tibia with patella-tibial suture; in other respects tibiae of all legs have similar structure. On larval legs patella-tibial suture crosses inner side of tibia at middle or at distal part (Kluge et al. 2017: Fig.11–12, 64, 122; Kluge et al. 2018: Fig.13-15, 17-18, 93) (see Index of characters [1.2.18], Index of characters [2.2.78.1]).

(14) Outer side of tibia near base is crossed by a row of fine colorless setae: portion of this row on posterior side of tibia is transverse, portion of this row on anterior side of tibia is oblique, on middle and hind tibia stretching along patella-tibial suture (Kluge et al. 2017: Fig.13-14; Kluge et al. 2018: Fig.17-18); such setae sparsely and irregularly arranged along posterior side of tibia and tarsus of all legs. The same in many other taxa.

(15) Small, arched setae, which are dispersed over larval leg, are especially dense on apex of inner (ventral) side of tibia, forming here a setal patch. Size of this setal patch varies individually; it is either limited by apical portion of tibia (Kluge et al. 2017: Fig.16–17), or is stretched along tibia. The same in Dabulamanzia and some other taxa.

(16) In subimago, all tarsal segments of all legs of both sexes are covered by pointed microlepides. The same in most Baetidae other than Baetungulata.

(17) Hind wing present in both sexes, of «Centroptilum-type»: narrow, with 2 longitudinal veins, with hooked costal projection (Kluge et al. 2017: Fig.51–55; Kluge et al. 2018: Fig.52, 106). Correspondingly, larva of both sexes have hind protoptera (Kluge et al 2018: Fig.14, 83).

(18) Larval caudalii of the «Siphlonuroid-type»: cerci and paracercus have equal length and bear long, dense primary swimming setae; in distal half of cercus outer side with more or less developed secondary swimming setae, not forming regular row (Kluge et al. 2017: Fig.63; Kluge et al. 2018: Fig.10).

(19) Before molt from larva to subimago, subimaginal gonostyli are bent under larval cuticle in «Cloeon-type» pose, i.e. with 2nd segments directed laterally (Kluge et al. 2017: Fig. 104; Kluge et al. 2018: Fig.57, 97). The same in most other Protopatellata and Anteropatellata-non-Baetovectata (see Index of characters [2.3.9]). 

() Fore wings with no more than one marginal intercalary in each space (unlike Baetovectata) [see Liberevenata (2)] (Index of characters [2.2.55]).