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

42 Bulletin 30 NOVEMBER 1, 2012breeding males with orange spots on the anal fin; breedingmales with quadrate lateral blotches (parallelismwith E. clinton); breeding males with no blue pigment onthe lips (reversal); and loss of palatine teeth (parallelismwith E. akatulo). Within this clade, E. jimmycarter and E.teddyroosevelt are united by four features of nuptial malecoloration, all reversals, including black marginal andsubmedial bands in the spinous dorsal fin, no blue pigmentin the soft dorsal fin, a ventrally-developed bluegreenbasicaudal bar, and no blue pigment in the analfin. Etheostoma teddyroosevelt exhibits two reversals as autapomorphies,loss of orange pigment on the face and lossof orange pigment in the base of the spinous dorsal fin.Etheostoma obama is characterized by three autapomorphiccharacter states: restriction of orange spots in the anal finof breeding males to the posteromedial portion of the fin,development of partly scaled cheeks, and polymorphismin the occurrence of palatine teeth.The relationships of E. obama, E. gore, and the E. jimmycarter-E.teddyroosevelt clade are unresolved in the minimum-lengthtree presented in Figure 25. In one of theother two minimum-length trees, a sister relationship betweenE. obama and E. gore is supported by the developmentof orange pigment on the face, which is interpretedas a parallelism with the same transformation in E. jimmycarter.However, this character state reconstruction is ambiguous,occurring only under DELTRAN optimization.The third minimum length tree places E. gore as sister tothe E. jimmycarter-E. teddyroosevelt clade. This relationshipis supported by the more extensive development of orangespots in the anal fin of nuptial males; transformation tothis state occurs under both optimization schemes butfrom different ancestral states. An E. gore-E. jimmycarter-E.teddyroosevelt clade is also supported by the derivation ofnaked cheeks, a parallel reversal with that in E. meadiae,but this occurs only under ACCTRAN optimization. UnderDELTRAN optimization the clade is also supportedby the absence of palatine teeth; this transformation shiftsdown the tree to the ancestor of the E. obama-E. gore-E.jimmycarter-E. teddyroosevelt clade under ACCTRAN. Additionalcharacters are clearly needed to more confidentlyresolve the relationships of these species.Evaluation of support.—The degree of support for thehypothesis of phylogenetic relationships (Fig. 25) was furtherevaluated by generating the strict and majority ruleconsensus trees for the minimum-length trees (60 steps)plus all trees that are one step longer. Twenty trees