Relationships of the Frogs of the Genus Ceratophrys (Leptodactylidae) and Their Bearing on Hypotheses of Pleistocene Forest Refugia in South America and Punctuated Equilibria

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Abstract
Of the 8 currently recognized species of Ceratophrys (C. aurita, C. calcarata, S. cornuta, C. cranwelli, C. ornata, C. pierottii, C. stolzmanni and C. testudo) 6 are considered to be species. C. testudo is considered a synonym of C. cornuta. The subfamily Ceratophryinae is argued to be monophyletic and is defined on the basis of 2 synapomorphies. The genus Ceratophrys (excluding C. pierottii) is monophyletic and is defined on the basis of 3 synapomorphies. Two subgenera are recognized within Ceratophrys. The subgenus Ceratophrys (1 synapomorphy) includes C. aurita, C. cranwelli and C. ornata where C. aurita and C. ornata are more closely interrelated. The subgenus Strombus (3 synapomorphies) includes C. calcarata, C. cornuta and C. stolzmanni where C. calcarata and C. cornuta are nearest relatives. C. pierottii is probably an intergeneric hybrid between C. cranwelli and Lepidobatrachus ilanensis. The least refuted hypothesis of relationships is inconsistent with the hypothesis that speciation among forest-dwelling species was mediated by Pleistocene contractions of forest into refugia. The northern species of Ceratophrys form a group fragmented in part by the Andes and the southern (and eastern) species form a 2nd group fragmented ecologically but not physically. The widespread Amazonian species, C. cornuta, shows little evidence of having had its distribution fragmented by forest contractions. C. cornuta is the most widely distributed species, is centrally distributed, and is by far the most apomorphous (measured in terms of autapomorphies). Such a combination is contrary to the expectations under the macroevolutionary hypothesis termed punctuated equilibria.

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