Title: Deiodination and the integration of larval adaptations in the Ceratophryinae (Anura: Leptodactylidae)
Description: As adults, ceratophryine frogs are characterized by hyperossified skulls, an enlarged gape, and an aggressive predatory nature. As larvae, however, members of the groups are truly remarkable, exhibiting an ecological and morphological diversity unmatched among anuran tadpoles. This diversity reflects the evolution of a derived feeding mode in two of the three genera that comprise the Ceratophryinae, larval carnivory. This thesis focuses on the process that reintegrates this larval diversity into a relatively conserved adult form--amphibian metamorphosis. To examine this question I employ an integrative approach, using phylogeny, morphology, gene expression, biochemistry, and primary organ culture. First, I use DNA sequence data to resolve the conflicting views of the evolutionary relationships among the Ceratophryinae and to examine the evolution of larval carnivory in the group. This analysis reveals that larval carnivory most likely arose twice independently, and that the evolutionary history of the group correlates with adaptation to an extreme environment--the and Chaco of South America. Next, I compare the metamorphic ontogeny of the larval jaws of representatives of the three genera that occur together in the Chaco, Chacophrys pierottii, Ceratophrys cranwelli , and Lepidobatrachus laevis . The results demonstrate the modular nature of anuran metamorphosis and the importance of overall larval period on the progression of metamorphic events.

To explore the local control of metamorphic timing in the jaws of these three species further, I use a biochemical assay to determine the ontogenetic profiles of type II 5'-deiodinase and type III 5-deiodinase in the lower jaw cartilages. The results indicate distinct roles for each enzyme in remodeling the jaws of ceratophryine frogs. To examine the roles of the deiodinases further, I use experimental manipulations of their activities on L. laevis lower jaw cartilages grown in culture. These experiments show that deiodinase activity in the lower jaw cartilages responds to thyroid hormone to regulate the initiation of metamorphic remodeling. Overall, this research demonstrates the utility of the integrative approach to the study of biological phenomena. In this case, using a natural experiment created by the evolutionary process to dissect the endocrine control of development in anuran amphibians.