(Percidae: Etheostoma), with Descriptions of Five New Species

40 Bulletin 30 NOVEMBER 1, 2012PHYLOGENETIC RELATIONSHIPSPhylogenetic hypothesisPhylogenetic analyses using both the unordered andmixed unordered-ordered character sets produced thesame three minimum-length trees with a length of 60steps and a consistency index of 0.683 (CI; Kluge andFarris, 1969) (uninformative characters excluded). Theretention index was 0.776 in the unordered analysis and0.787 in the mixed analysis (RI; Farris, 1989). The leastresolved of the minimum-length trees, which also representsthe strict consensus tree, is shown in Figure 25.All three trees support the monophyly of Doration with E.akatulo sister to a clade containing all other species in thesubgenus. A pattern of asymmetrical branching proceedsup the tree with E. clinton, E. stigmaeum, E. meadiae, andE. jessiae. Etheostoma jessiae is sister to a clade containing E.obama, E gore, E. jimmycarter, and E. teddyroosevelt. The othertwo minimum-length trees differ only in the resolution ofthe trichotomy involving this latter clade. One tree placesE. gore as sister to the E. jimmycarter-E. teddyroosevelt clade,and the other tree unites E. obama and E. gore as sister tothe E. jimmycarter-E. teddyroosevelt clade. As the more conservativeestimate of relationships, the less highly resolvedtree is presented as the preferred phylogenetic hypothesis(Fig. 25).The monophyly of Doration is supported by ten derivedcharacter states (Fig. 25). Breeding males of the ancestorof this clade are hypothesized to have possessed a seriesof 7–11 vertically elongate blue bars, scattered red-orangemarkings on the sides, iridescent blue or blue-green pigmenton the opercles, cheeks, lips, and mid-gular area, aspinous dorsal fin with dusky to black marginal and submedialbands and red-orange medial band, and a ventrally-developedblue-green basicaudal bar. The ancestor alsopossessed a pair of small dark spots on the medial caudalfin base and modally 11–13 dorsal spines.Within Doration, E. akatulo is sister to a clade containingall other species in the subgenus. This latter clade is unitedby three derived features of male breeding coloration:blue or blue-green marginal and submedial bands in thespinous dorsal fin, blue or blue-green pigment in the baseof the soft dorsal fin, and blue or blue-green pigment inthe base of the anal fin. Etheostoma akatulo is characterizedby three homoplasious autapomorphies: a complete lateralline, fully scaled cheeks, and loss of palatine teeth. Thecomplete lateral line is interpreted in this optimizationas a reversal, but an equally parsimonious reconstructionallows for parallel derivation of an incomplete lateral linein E. caeruleum, E. chlorosoma, and the ancestor of the cladesister to E. akatulo. Derivation of fully scaled cheeks in E.akatulo is an instance of parallel evolution of the samestate in the ancestor of the E. obama-E. gore-E. jimmycarter-E. teddyroosevelt clade, and the loss of palatine teeth is aparallel transformation with that in E. chlorosoma.Monophyly of the clade containing E. stigmaeum, E.meadiae, E. jessiae, E. obama, E. gore, E. jimmycarter, and E.Figure 25. Hypothesized phylogeny of species of thesubgenus Doration. This is one of three minimum-lengthtrees and also represents the strict consensus tree.Derived character states supporting each of the letteredbranches are: A) 17(1), 19(1), 22(1), 34(1); B) 16(1), 25(1),26(1), 30(1), 34(2); C) 17(0)*, 23(0)*, 27(1)*; D) 28(0)*,29(0)*; E) 1(1), 2(1), 3(1), 4(1), 5(1), 7(1), 8(1), 12(1), 20(1),32(1); F) 23(0)*, 24(1)*, 28(0)*; G) 7(2), 10(1), 13(1); H)1(2)*, 27(1)*; I) 12(2); J) 28(1)*; K) 5(0)*, 8(2); L) 22(0,1)*,24(0,1)*; M) 9(1), 11(1), 15(1); N) 22(0)*, 28(1)*; O)1(2)*, 4(0)*, 6(1), 14(2), 24(1)*; P) 14(1), 28(1)*; Q) 7(1)*,10(0)*, 12(1)*, 13(0)*; R) 6(0)*, 9(0)*. See Appendix A forcharacter descriptions and Appendix B for data matrix.Homoplasious transformations indicated by asterisks.teddyroosevelt is supported by a single character, a verticalblue or blue-green basicaudal bar in nuptial males that extendsfrom the dorsal to the ventral margin of the caudalfin. Etheostoma clinton, sister to this clade, exhibits quadratelateral blotches in breeding males and 9 preoperculomandibularpores as autapomorphies. Both states representinstances of parallel development, the former withthe ancestor of the E. obama-E. gore-E. jimmycarter-E. teddyrooseveltclade, and the latter with E. nigrum.The clade containing E. meadiae, E. jessiae, E. obama,E. gore, E. jimmycarter, and E. teddyroosevelt is united by theabsence of blue or blue-green pigment on the gular area