Anuran amphibians undergo major morphological transitions during development, but the contribution of their markedly different life-history phases to macroevolution has rarely been analysed. Here we generate testable predictions for coupling versus uncoupling of phenotypic evolution of tadpole and adult life-history phases, and for the underlying expression of genes related to morphological feature formation. We test these predictions by combining evidence from gene expression in two distantly related frogs, Xenopus laevis and Mantidactylus betsileanus, with patterns of morphological evolution in the entire radiation of Madagascan mantellid frogs. Genes linked to morphological structure formation are expressed in a highly phase-specific pattern, suggesting uncoupling of phenotypic evolution across life-history phases. This gene expression pattern agrees with uncoupled rates of trait evolution among life-history phases in the mantellids, which we show to have undergone an adaptive radiation. Our results validate a prevalence of uncoupling in the evolution of tadpole and adult phenotypes of frogs.

Many tropical rainforests harbour speciesrich assemblages of frogs and, consequently, of tadpoles. These larvae are often morphologically highly diverse, especially in their oral structures. Whether this might represent an important axis of ecological partitioning in frog assemblages remains an untested hypothesis. In general, it is poorly known how diverse tadpole assemblages are organised. Using information from Madagascar's remarkably species-rich stream tadpole assemblages, we analysed the distribution and co-occurrence of 44 species. We also assessed the importance of phylogenetic history and eco-morphological adaptation for habitat selection at two functional and two spatial levels. We show that both rare and common species exist in most morphological clusters and phylogenetic groups of tadpoles. Habitat characteristics of the streams and surrounding forest influence species composition. Whereas there is a general trend in preferring wide and deep streams without a steep slope for most species, some of the morphological clusters separate along specific habitat variables. Stream choice is influenced both by phylogenetic history and morphological adaptation. Within streams, tadpoles partition microhabitat mainly according to their morphological cluster but without phylogenetic signal. Species dissimilarities of habitat selection on both spatial scales are not correlated. We found no evidence for competition in the tadpole assemblages studied.

Frogs in the genus Gephyromantis from Madagascar were assumed to have a direct developmental mode, i.e. the complete embryonic and larval development within the egg, but recently free-swimming, exotrophic tadpoles of a few species have been found. Herein we provide detailed morphological descriptions of the tadpoles of five more species of this genus, indicating a developmental mode other than direct development in species of Gephyromantis. Tadpoles of Gephyromantis granulatus, G. sculpturatus, G. tschenki, and G. ventrimaculatus were found free-swimming in streams, and tadpoles of G. sp. aff. blanci were raised after hatching from clutches found in the leaf litter. All tadpoles were identified by DNA barcoding. The oral discs of all five species are characterized by the lack of many typical morphological traits of exotrophic tadpoles (such as oral papillae and keratodonts). This indicates that these tadpoles are either non-feeding (endotrophic) or only facultatively feeding tadpoles. We classify these tadpoles as nidicolous based on the observation that the larvae of G. sp. aff. blanci stayed after hatching in the jelly nest until metamorphosis. It remains unclarified whether all species have strictly nidicolous tadpoles, and the larvae of the four species found in the streams were just accidentally washed into these streams; alternatively, some of these tadpoles might be nidicolous at first but in some species need to live in free water at later stages.