Phylogenetic relationships of bufonid frogs and tests of alternate macroevolutionary hypotheses characterizing their radiation

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Abstract
The frog family Bufonidae is a large group distributed throughout all major land masses of the world except the polar regions. Although the monophyly of Bufonidae is strongly supported, phylogenetic relationships within the group are not well understood. Because of apparently large differences in rates of morphological and behavioral evolution, speciation, and geographical range, Bufonidae raises intriguing macroevolutionary questions. For example, one might imagine that the group's evolutionary history is congruent (1) with its morphological and behavioral character distribution, or (2) with its current geographical positioning. To examine phylogenetic relationships within Bufonidae and test these alternate hypotheses, DNA sequence data were obtained from four genes, three mitochondrial and one nuclear, from populations throughout the geographic and taxonomic breadth of the family. Phylogenetic analyses of these sequences by several methods support several hypotheses of relationships, notably: (1) contrary to the previous hypotheses of some workers, Bufo is not the basal, wildly paraphyletic genus of Bufonidae, although it is also not monophyletic, (2) the basal splits within bufonids separate South American «atelopodid» genera from all other bufonids, although the monophyly of the «atelopodids» is not supported, and (3) the most highly supported clades within the family generally correspond to small- to medium-sized, geographically and morphologically homogeneous groups. These trees are used to test the two alternate macroevolutionary hypotheses, and, while neither hypothesis fully explains the observed phylogenetic relationships, these relationships are broadly consistent with both character distribution and geography at particular levels.

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