Historical biogeography and a mitochondrial DNA phylogeny of grouse and ptarmigan.

We sequenced 2690 nucleotides of mitochondrial DNA (mtDNA) including the entire control region (CR), partial 12S and 16S ribosomal RNAs, NADH dehydrogenase subunit 2, and cytochrome b genes from representatives of all the 17 living species of grouse and ptarmigan (Aves; Galliformes; subfamily Tetraoninae). Substitution rates and phylogenetic signals were variable among genes, with the CR being more informative than protein-coding and rRNA genes. Phylogenetic trees, computed with the CR or the concatenated sequences, indicate that: (1) genus Bonasa is monophyletic and basal within the subfamily, (2) all the other currently recognized genera of Tetraoninae are monophyletic, except Dendragapus; (3) D. obscurus is related to Centrocercus urophasianus and divergent from former D. canadensis and D. falcipennis, which, accordingly, may be ascribed to the distinct genus Falcipennis; (4) Tympanuchus, Dendragapus, and Centrocercus form a clade comprising taxa distributed exclusively in North America; and (5) the North American species of Bonasa (B. umbellus) and Lagopus (L. leucurus) are basal to their Eurasian and Holarctic congeneric species. These findings, and a dispersal-vicariance analysis, support a North American origin of the subfamily and of all the genera of Tetraoninae, with the possible exception of Tetrao. Present species distributions might have been attained by at least three dispersal events from North America to Eurasia, involving the ancestors to Palearctic Bonasa, the ancestors to circumpolar Lagopus mutus/L. lagopus, and the clade leading to Tetrao/Falcipennis. According to a "standard calibration" of the mtDNA molecular clock (2% sequence divergence per million years), Bonasa split about 5-6 million years ago, the other genera diverged during the upper Pliocene, and most of the congeneric species with North American and Eurasian distributions (Bonasa, Lagopus, and Falcipennis) originated during the lower Pleistocene, well before the last interruption of the Beringian land bridge.