Chaoborus flavicans species complex We found evidence for at least four species in the Chaoborus flavicans complex: Chaoborus flavicans, C. albipes, C. posio sp. n. and a lineage from Japan. The monophyly for each of these species is strongly supported by bootstrap values (Fig. 15) and is not attributable to random branching under a coalescent null model (Rosenberg’s P(AB) 30%) indicates that divergence of the complex likely occurred well before the Pleistocene. Moreover, widespread species such as C. albipes and C. flavicans with two or more closely related geographic clades may also have been present before the Pleistocene. Chaoborus tertiarius (von Heyden), Upper Oligocene fossil species, is composed of two complete pupae, disarticulated pupal parts, larval mandibles, and anal fans (Borkent 1978). This taxon may be ancestral to C. (Chaoborus s.str.) and the subgenus C. (Schadonophasma Dyar & Shannon 1924), or it may be ancestral, or a sister, to species of C. flavicans species complex (Borkent 1978). Among the fossil material are two types of larval mandibles: the subordinate tooth is either present between teeth two and four or it branches from the second tooth (Borkent 1978, figs. 3 A-H). However, Borkent admits that the subordinate tooth of the flavicans - type “was very difficult to see”, and hence it is uncertain if C. tertiarius is closely related to the C. flavicans species complex. Diagnosis. Adults male. Penultimate flagellomere slightly longer than ultimate, or about equal in length. Flagellomeres pale or partly darkened, bases of whorls dark, giving annulated appearance. Scutellar stripes bare, orange brown–almost black, pleural sclerites pale yellow and with a varying degree of darker coloration. Legs straw yellow or grayish, apical tarsomeres may be somewhat darkened. Tergites yellowish brown or dark brown, bases of setae with a dark ring. Male epandrium either triangular or broadly rounded in shape, its length about 0.35–0.37 times the length of gonocoxite. Male paramere with a conspicuous apical claw, elongated, almost straight or curved. Pupa. Outer rib of terminal process smooth, apical spines absent. Mid rib of terminal process usually darker than lateral ribs, margin of inner rib serrated along apical 2/3. Length/width ratio of 8 th segment ca. 0.36. Respiratory organ widest medially, constricted apically or not. IV instar larva. Total length 7–15 mm (e.g. Parma 1971b). Labral blade elongated, degree of serration varied. Mandibular subordinate tooth (3) positioned at the vertex of tooth 2 and tooth 4. Mandibular lateral teeth either small and inconspicuous or larger and distinguishable. Dorsal process pointed. Key to species of Chaoborus flavicans species complex Note. All species key out as C. flavicans in the key by Saether (1972). adult males 1 Gonocoxite and gonostylus stout in structure, length:width ratio 2.64 (2.4–3) and 9.43 (8–10.7), respectively (Fig. 14a); apical claw of paramere relatively long (Fig. 14 c–e).............................................. C. posio Salmela sp. n. - Gonocoxite and gonostylus slender, length:width ratio ca. 3–3.5 (2.37–3.9) and 12.5–13.7 (10.2–16), respectively (Figs. 6a, 11a); apical claw of paramere relatively short (Figs. 6 c–e, 11c–e)................................................ 2 2 Paramere medially bent and constricted; apical claw rather narrow, moderately curved (Figs. 6, 8a).... C. flavicans (Meigen) - Paramere medially almost straight or gently curved; apical claw stout, curved (Figs. 8b, 11)...................................................................................................... C. albipes (Johannsen stat. rev.) pupaeSeparation of pond-dwellingC. flavicansandC. albipespupae may be obscured by intraspecific variation 1 Respiratory organ club-shaped, lacking subapical constriction, relatively short (844 (770–930) µm, Fig. 9d................................................................................................ C. posio Salmela sp. n. - Respiratory organ with subapical constriction, either slender or voluminous (Figs. 9 a–c)............................. 2 2 Length of respiratory horn Ẑ 1000 µm (920–1360), either slender or voluminous (Figs. 9 a–b)......... C. flavicans (Meigen) - Length of respiratory organ mostly 16........................................................................... 2 2 Number of mandibular fan bristles>20, usually>22 (21–29) (Fig. 10h); labral blade finely serrated, rather wide (length:width ratio 3,7 (3–5 Fig. 10g)............................................................... C. posio Salmela sp. n. - Number of mandibular fan bristles up to 25, usually 15–21; labral blade almost devoid of serration, slender (length:width ratio 5.59 (4.38–6.85, Fig. 10e)..................................................... C. albipes (Johannsen, stat. rev.)

Published as part of Salmela, Jukka, Härmä, Oskari & Taylor, Derek J., 2021, Chaoborus flavicans Meigen (Diptera, Chaoboridae) is a complex of lake and pond dwelling species: a revision, pp. 151-196 in Zootaxa 4927 (2) on pages 154-155, DOI: 10.11646/zootaxa.4927.2.1, http://zenodo.org/record/4536900