Red- and yellow-footed tortoises, Chelonoidis carbonaria and C. denticulata (Reptilia: Testudines: Testudinidae), in South American savannahs and forests: do their phylogeographies reflect distinct habitats?

Using sequence data of the mitochondrial cytochrome b gene, we investigated phylogeographic differentiation of the Amazonian tortoise species Chelonoidis carbonaria and C. denticulata. While C. carbonaria is generally restricted to savannah habitats and adjacent forests, C. denticulata is associated with wet tropical and subtropical forests. Our study suggests a correlation between distinct habitat preferences and phylogeography of the two species. In Maximum Parsimony, Maximum Likelihood and Bayesian analyses, haplotypes of C. carbonaria cluster in several distinct clades reflecting the species' patchy distribution in savannah habitats. By contrast, haplotypes of C. denticulata are only weakly differentiated; a finding also confirmed by parsimony network analysis. This suggests that the contiguous Amazonian rainforest allows gene flow between populations of the forest-dwelling C. denticulata throughout the range, but significantly impedes gene flow in C. carbonaria. The phylogeographic structure and extant distribution pattern of C. carbonaria is supportive of former Amazonian rainforest fragmentation, enabling the dispersal of savannah species. Based on fossil calibration, we dated divergence times for the C. carbonaria clades using a relaxed molecular clock, resulting in average estimates ranging from 4.0–2.2 mya. This implies that the onset of rainforest fragmentation could predate the Pleistocene considerably. Furthermore, our findings call for further research on geographic and taxonomic variation in C. carbonaria and for a reassessment of the conservation status of the distinct genetic units.