The Australoheros (Teleostei: Cichlidae) species of the Uruguay and Paraná River drainages

" Australoheros Rícan and Kullander, 2006 includes four described species Australoheros facetus (Jenyns, 1842), Australoheros tembe (Casciotta, Gómez & Toresani, 1995), Australoheros scitulus (Rícan & Kullander, 2003) and Australoheros kaaygua Casciotta, Almirón & Gómez, 2006. Four additional species are newly described in this paper based on results presented in Rícan and Kullander (2006): Australoheros forquilha sp. nov., from the tributaries of the Upper Río Uruguay in Brazil; Australoheros charrua sp. nov. and Australoheros minuano sp. nov., from the tributaries of the Middle Río Uruguay in Brazil and Uruguay, and Australoheros guarani sp. nov. from the Río Paraná tributaries in Paraguay. Sympatric species are not found closely related and all sister species have allopatric distributions. Four species groups are recognized based on phylogenetic hypotheses generated from morphological and cyt b characters; the forquilha group; A. forquilha, A. tembe; the scitulus group: A. scitulus, A. charrua; the kaaygua group: A. kaaygua, A. minuano; and the facetus group: A. facetus, A. guarani "