Phylogeny, Molecules Versus Morphology, and Rates of Character Evolution Among Fruitbats (Chiroptera: Megachiroptera)

Abstract

Andersen's 1912 monograph on megachiropterans remains the definitive work on the systematics of this group. Andersen argued that the Macroglossinae, containing the eonycterine and notopterine sections, are a monophyletic sister-group to other fruitbats (i.e. Andersen's Rousettus, Cynopterus and Epomophorus sections). Two recent molecular studies (DNA hybridisation and restriction mapping of ribosomal cistrons), as well as an analysis of female reproductive characters, challenge the monophyly of the Macroglossinae and several of Andersen's other conclusions such as the phylogenetic position of Nyctimene. We performed a cladistic analysis on 36 morphological characters, including 33 that were gleaned from Andersen, to determine whether phylogenetic hypotheses based on modem phylogenetic methods are in agreement with Andersen's original conclusions and to compare morphological and molecular phylogenetic hypotheses. Minimum-length trees based on parsimony are largely consistent with Andersen and support (1) a monophyletic Macroglossinae, within which the eonycterine section is paraphyletic with respect to a monophyletic notopterine section, (2) a monophyletic Cynopterus section, excepting the exclusion of Myonycteris, (3) a monophyletic Epomophorus section, excepting the exclusion of Plerotes, and (4) a paraphyletic Rousettus section, with several of the Rousettus-like forms branching off near the base of the tree. Bootstrapping analyses on a reduced data-set that included taxa shared in common with the DNA hybridisation study did not provide strong support (greater than or equal to 95%) for any clades but did provide moderate support (greater than or equal to 70) for several clades, including a monophyletic Macroglossinae. These findings are in marked contrast to the DNA hybridisation phylogeny. A high index of between-data-set incongruence is further evidence for the clash between DNA hybridisation and morphology. A phylogenetic framework was constructed on the basis of morphological data and DNA hybridisation data using a criterion of moderate support and shows little resolution, whereas employing a criterion of strong support produced a framework resolving several additional nodes. One implication of this framework is that characteristic macroglossine features such as a long tongue with a thick carpet of filiform papillae have evolved independently on several occasions (or evolved once and were lost several times). Rates of character evolution for the morphological characters employed in our analysis were calculated using divergence times estimated from DNA hybridisation data. Rates have apparently been fastest in the interior branches, and slower along the external branches, which suggests an early adaptive radiation in the history of fruitbats.