Philippine bent-toed geckos of the Cyrtodactylus agusanensis complex: phylogeny, morphological diversity, and description of three new species

We review the taxonomic status of Philippine bent-toed geckos previously referred to Cyrtodactylus agusanensis. We delineate four evolutionary lineages within the C. agusanensis complex from the southeastern islands of the archipelago and describe three of these lineages as new species. The new species and true C. agusanensis are identified by numerous, nonoverlapping morphological characters and by allopatric ranges on separate islands. Our morphology-based taxonomic conclusions are bolstered by biogeographic evidence and marked interspecific divergence between monophyletic groups defined by mitochondrial and nuclear DNA sequences. To compliment these descriptions and enable future taxonomic work on Philippine Cyrtodactylus, we rediagnose and redescribe C. agusanensis. Because the holotype of C. agusanensis was destroyed in World War II, we designate a neotype for this species and restrict its geographic range to north central Mindanao Island. Our phylogenetic estimate suggests that the C. agusanensis complex originated in Mindanao and spread progressively north, diversifying incrementally with colonization of successive islands in a south-to-north pattern of biogeographic expansion and allopatric speciation.