Taxonomic revision of the genus Pungitius with emphasis on P. hellenicus

Variation in morphology of species of the genus Pungitius is enormous. Due to this variation, the taxonomic status of many taxa is uncertain; while some workers recognize two species, P. pungitius and P. platygaster, others recognize more. I examined the seven nominal species (P. pungitius, P. occidentalis, P. sinensis, P. tymensis, P. platygaster, P. laevis and P. hellenicus) to determine their taxonomic status, especially the validity of P. hellenicus from Greece, and to hypothesize their systematic relationships within the genus. Twenty four traditional morphometric characters and 21"truss distances". were measured and fifteen meristic characters were counted on 310 specimens from much of the range of the species. The osteology of P. hellenicus was compared to that of the other species. Morphometric and meristic characters were of little value in differentiating among the species because of the variation and overlap of their values. However, I recognize three valid species in the genus, namely, P. hellenicus, P. platygaster, and P. pungitius with five subspecies (P. p. pungitius, P. p. laevis, P. p. tymensis, P. p. sinensis, and P. p. occidentalis). Pungitius hellenicus can be recognized by a combination of five characters: fewer than seven dorsal spines,"reduced ectocoracoid"., absence of a keel on the caudal peduncle, and lack of the pelvic girdle and large lateral scutes. The phylogenetic relationship of the species is uncertain, but it seems that P. hellenicus is the most primitive sister group of the other species. However, P. hellenicus may be a neotenic form of P. platygaster. The phylogenetic relationships of Gasterosteus, Culaea, and Pungitius are also uncertain; however, this study suggests that Culaea inconstans is a member of the genus Pungitius.