The anuran Bauplan: a review of the adaptive, developmental, and genetic underpinnings of frog and tadpole morphology.

Anurans (frogs, toads, and their larvae) are among the most morphologically derived of vertebrates. While tightly conserved across the order, the anuran Bauplan (body plan) diverges widely from that of other vertebrates, particularly with respect to the skeleton. Here we address the adaptive, ontogenetic, and genetic bases of three such hallmark anuran features: (1) the absence of discrete caudal vertebrae, (2) a truncated axial skeleton, and (3) elongate hind limbs. We review the functional significance of each as it relates to the anuran lifestyle, which includes locomotor adaptations to both aquatic and terrestrial environments. We then shift our focus to the proximal origins of each feature, namely, ontogeny and its molecular regulation. Drawing on relatively limited data, we detail the development of each character and then, by extrapolating from comparative vertebrate data, propose molecular bases for these processes. Cast in this light, the divergent morphology of anurans emerges as a product of evolutionary modulation of the generalised vertebrate developmental machinery. Specifically, we hypothesise that: (1) the formation of caudal vertebrae is precluded due to a failure of sclerotomes to form cartilaginous condensations, perhaps resulting from altered expression of a suite of genes, including Pax1, Pax9, Msx1, Uncx-4.1, Sonic hedgehog, and noggin; (2) anteriorised Hox gene expression in the paraxial mesoderm has led to a rostral shift of morphological boundaries of the vertebral column; and, (3) spatial and temporal shifts in Hox expression may underlie the expanded tarsal elements of the anuran hind limb. Technology is currently in place to investigate each of these scenarios in the African clawed frog Xenopus. Experimental corroboration will further our understanding of the molecular regulation of the anuran Bauplan and provide insight into the origin of vertebrate morphological diversity as well as the role of development in evolution.