Morphological Variation in Anuran Limbs: Constraints and Novelties.

Anurans have three primary types of locomotion: walking, jumping, and swimming. Additionally, they may dig, climb, grasp, etc. All adult anurans have four limbs, with four fingers on the hands and five toes on the feet. We summarized and updated knowledge on the interspecific variation within anuran limbs, then discuss how developmental constraints (e.g., in size) and novelties may have influenced anuran diversification through the locomotion. We analyze morphological variation from limb bud stages up to the final limb form resulting from certain skeletal organization and growth. We find limited morphometric variations in the skeleton of different developmental modules (i.e., skull, trunk, urostyle, limbs) indicate that the anuran body shape is largely constrained. We identify specializations of the stylopodium, zeugopodium, and proximal carpals/tarsals that have evolved to facilitiate saltatorial locomotion. We show that the anuran prepollex and prehallux are not vestigial digits and that they have come to serve specialized function. Medial rotation of the manus in anurans appears to have evolved to help distribute the force of impact upon landing at the end of a jump. Additional skeletal elements in anuran limbs are intercalary elements and sesamoids. The intercalary elements appear within neobatrachians and are integrated with digital pads in lineages capable of locomotion on smooth vertical surfaces. They have allowed arboreal anurans to occupy a wide range of arboreal habitats.