The arboreal frogs in the genus Spinomantis (Amphibia: Mantellidae) from Madagascar: a survey of molecular, morphological and bioacoustic variation, with descriptions of four new species

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We review the systematics of the Madagascar-endemic frogs of the genus Spinomantis based on DNA sequences of mitochondrial and nuclear-encoded genes from 368 individuals. We find the arboreal species in the genus, the focus of this study, to form a well-supported clade in a mitochondrial multi-gene tree, containing three subclades, here named the peraccae clade, the aglavei clade, and the massi clade. Each of these contained numerous deep mitochondrial lineages differing by at least 3.1% uncorrected pairwise distance in a fragment of the mitochondrial 16S rRNA gene. Our preferred partition distinguishes 20 mitochondrial species-level lineages in the arboreal Spinomantis clade, including seven nomi­nal species, four new species described herein, and nine additional candidate species that require further study. The four new species described herein are delimited by a combination of several lines of evidence, including high divergence in mitochondrial DNA (> 3.5% 16S divergence to nominal species), isolated phylogenetic position, differentiation in nuclear-encoded gene fragments, and morphological and bioacoustic divergence: S. chlorostictus sp. n. is a species of the peraccae clade primarily from northern Madagascar differing by color pattern and bioacoustics from S. peraccae; S. royae sp. n. and S. leonieae sp. n., belonging to the aglavei clade, are morphologically similar to S. fimbriatus but differ by color pattern and advertisement call; and S. fohitsilo sp. n. is morphologically similar to S. massi but not that species’ direct sister group, and differs by its advertisement call. Our study provides a baseline for understanding the diversity and biogeography of Spinomantis, highlights the enormous amount of yet unexplored amphibian diversity in Madagascar, and points to geo­graphical regions and taxa where further field exploration, including the collection of voucher specimens and of bioacous­tic data, is needed to complete the herpetofaunal inventory of the island.

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