Old World fruitbat phylogeny: Evidence for convergent evolution and an endemic African clade

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Abstract
Knud Andersen (1912, Catalogue of the Chiroptera in the Collections of the British Museum: I. Megachiroptera, British Museum of Natural History, London) divided Old World fruitbats (family Pteropodidae) into the rousettine, cynopterine, epomophorine, eonycterine, and notopterine sections. The latter two sections comprise the subfamily Macroglossinae; members of this subfamily exhibit specializations for nectarivory (e.g., elongated, protrusible, brushy tongues) and cluster together in cladistic analyses based on anatomical characters. Other evidence, including single-copy DNA hybridization, suggests that macroglossines are either paraphyletic or polyphyletic; this implies that adaptations for pollen and nectar feeding evolved independently in different macroglossine lineages or were lost in nonmacroglossines after evolving in a more basal common ancestor. Hybridization data also contradict Andersen’s phylogeny in providing support for an endemic African clade that includes representatives of three of Andersen’s sections. Here, we present complete mitochondrial 12S rRNA and valine tRNA gene sequences for 20 pteropodids, including representatives of all of Andersen’s sections, and examine the aforementioned controversies. Maximum likelihood, minimum evolution, and maximum parsimony analyses all contradict macroglossine monophyly and provide support for an African clade that associates Megaloglossus and Lissonycteris and those two with Epomophorus. In conjunction with the DNA hybridization results, there are now independent lines of molecular evidence suggesting: (i) convergent evolution of specializations for nectarivory, at least in Megaloglossus versus other macroglossines, and (ii) a previously unrecognized clade of endemic Africa taxa. Estimates of divergence time based on 12S rRNA and DNA hybridization data are also in good agreement and suggest that extant fruitbats trace back to a common ancestor 25 million to 36 million years ago.

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