The members of the cyprinid subfamily Danioninae form a diverse and scientifically important group of fishes, which includes the zebrafish, Danio rerio. The diversity of this assemblage has attracted much scientific interest but its monophyly and the relationships among its members are poorly understood. The phylogenetic relationships of the Danioninae are examined herein using sequence data from mitochondrial cytochrome b, mitochondrial cytochrome c oxidase I, nuclear opsin, and nuclear recombination activating gene 1. A combined data matrix of 4117. bp for 270 taxa was compiled and analyzed. The resulting topology supports some conclusions drawn by recent studies on the group and certain portions of the traditional classification, but our results also contradict key aspects of the traditional classification. The subfamily Danioninae is not monophyletic, with putative members scattered throughout Cyprinidae. Therefore, we restrict Danioninae to the monophyletic group that includes the following genera: Amblypharyngodon, Barilius, Cabdio, Chela, Chelaethiops, Danio, Danionella, Devario (including Inlecypris), Esomus, Horadandia, Laubuca, Leptocypris, Luciosoma, Malayochela, Microdevario, Microrasbora, Nematabramis, Neobola, Opsaridium, Opsarius, Paedocypris, Pectenocypris, Raiamas, Rasbora (including Boraras and Trigonostigma), Rasboroides, Salmostoma, Securicula, and Sundadanio. This Danioninae sensu stricto is divided into three major lineages, the tribes Chedrini, Danionini, and Rasborini, where Chedrini is sister to a Danionini-Rasborini clade. Each of these tribes is monophyletic, following the restriction of Danioninae. The tribe Chedrini includes a clade of exclusively African species and contains several genera of uncertain monophyly (Opsarius, Raiamas, Salmostoma). Within the tribe Rasborini, the species-rich genus Rasbora is rendered non-monophyletic by the placement of two monophyletic genera, Boraras and Trigonostigma, hence we synonymize those two genera with Rasbora. In the tribe Danionini, the miniature genus Danionella is recovered as the sister group of Danio, with D. nigrofasciatus sister to D. rerio.
Bibliographical noteFunding Information:
This study was funded in part by the National Science Foundation (USA) Assembling the Tree of Life initiative Grants EF 0431326 (Mayden/Wood) and EF 0431132 (Simons) for the Cypriniformes Tree of Life project. We are indebted to K. Conway for his help on this project, from inception to completion, his contributions were most significant with identification of taxa and during the writing phase, with his knowledge of the literature and in discussions of nomenclature and taxonomy. We are grateful for the assistance and sequence data provided by W.-J. Chen, his involvement was key during the data collection phase, where his skills and expertise allowed for the successful completion of the data set used in this study. The authors wish to thank the following individuals and their affiliated institutions for providing specimens: P. Berendzen (Northern Iowa University); R. Britz (British Museum of Natural History); D. Catania (California Academy of Sciences); A. Golubtsov (Russian Academy of Sciences); B. Kuhajda (University of Alabama); T. Moritz (BMNH); D. Neely (CAS); F. Pezold (Texas A&M University—Corpus Christi); R. Schelly (American Museum of Natural History); M. Stiassny (AMNH); and L. Yang (Chinese Academy of Sciences/Saint Louis University). The manuscript benefited from discussions of nomenclature and taxonomy with C. Ferraris and D. Frost. We thank J. Baker for assistance with data analysis, C. Dillman for assistance with data output and tree drawing, L. Rüber for helpful comments on an earlier draft of this manuscript and for making available to us some sequence data, and L. Yang for providing and translating Chinese citations. The authors wish to acknowledge the Willi Hennig Society for making the TNT software freely available.
- Cytochrome b
- Cytochrome c oxidase I