The cyprinid tribe Labeonini (sensu Rainboth, 1991) is a large group of freshwater fishes containing around 40 genera and 400 species. They are characterized by an amazing diversity of modifications to their lips and associated structures. In this study, a total of 34 genera and 142 species of putative members of this tribe, which represent most of the generic diversity and more than one third of the species diversity of the group, were sampled and sequenced for four nuclear genes and five mitochondrial genes (totaling 9465. bp). Phylogenetic relationships and subdivision of this tribe were investigated and the placement and status of most genera are discussed. Partitioned maximum likelihood analyses were performed based on the nuclear dataset, mitochondrial dataset, combined dataset, and the dataset for each nuclear gene. Inclusion of the genera Paracrossochilus, Barbichthys, Thynnichthys, and Linichthys in the Labeonini was either confirmed or proposed for the first time. None of the genera Labeo, Garra, Bangana, Cirrhinus, and Crossocheilus are monophyletic. Taxonomic revisions of some genera were made: the generic names Gymnostomus Heckel, 1843, Ageneiogarra Garman, 1912 and Gonorhynchus McClelland, 1839 were revalidated; Akrokolioplax Zhang and Kottelat, 2006 becomes a junior synonym of Gonorhynchus; the species Osteochilus nashii was found to be a member of the barbin genus Osteochilichthys. Five historical hypotheses on the classification of the Labeonini were tested and rejected. We proposed to subdivide the tribe, which is strongly supported as monophyletic, into four subtribes: Labeoina, Garraina, Osteochilina, and Semilabeoina. The taxa included in each subtribe were listed and those taxa that need taxonomic revision were discussed.
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We would like to express our gratitude to Junbing Li, Xuzhen Wang, and Kevin Conway for their help with sampling. Paula Mabee is thanked for her help on transferring samples. The Natural History Museum of the University of Kansas and Andrew Bentley, and the Cornell University Museum of Vertebrates are thanked for loans of specimens. We are grateful to E Zhang, Lanping Zheng, Kevin Tang, and Susana Schönhuth for helpful discussion. Yahui Zhao and Yanfei Huang are thanked for providing information on Protolabeo and Stenorynchoacrum. We thank Hsin-Hui Wu for his help on using the Cluster computing system at Saint Louis University. We are also grateful to the associate editor and two anonymous reviewers for their very valuable comments and suggestions that helped improve this paper. This research was supported in part by personal funds of RLM, by the USA National Science Foundation grants including The Cypriniformes Tree of Life initiative (CToL) (EF 0431326 to RLM; EF 0431132 to AMS) and the Collaborative PBI All Cypriniformes Species – Phase II of An Inventory of the Otophysi (ACSI-II) (DEB1021840 to RLM), by the Japan Society for the Promotion of Science (17207007 and 22370035) and by Saint Louis University, St. Louis, Missouri.
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