Skull development during anuran metamorphosis: III. Role of thyroid hormone in chondrogenesis

Metamorphosis of cranial cartilages in anuran amphibians constitutes one of the most dramatic and extensive ontogenetic transformations in vertebrates. We quantitatively examined the role of thyroid hormone (3,3',5-triiodo-L-thyronine; T3) in mediating gross aspects of this morphological repatterning in the skull of the Oriental fire-bellied toad, Bombina orientalis. T3 was administered via plastic (Elvax) micropellets in three treatment dosages (2.5, 0.25, and 0.025 microgram) and one control dosage (0 microgram) to tadpoles of three Gosner developmental stages--28/29, 30/31, and 32/33; tadpoles were recoved up to 8 d (treatment and control dosages) or 14 d (control dosage) later. Response of larval cartilages to exogenous T3 was dosage dependent but not implant-stage dependent; chondrogenic tissues that participate in metamorphic transformation are competent to respond to T3 well before they normally do. Metamorphic effects of T3 were visible within 2 d; in most treatment groups, the normal metamorphic sequence was two-thirds complete after 8 d. While T3 also induced precocious ossification, the normal temporal relation between bone formation and cartilage transformation was dissociated in experimental groups. Morphological integration between cartilage and bone during cranial metamorphosis is at least partly the result of each tissue responding independently to endocrine factors.     

Patterns of reproductive‐mode evolution in Old World tree frogs (Anura,Rhacophoridae)

The Old World tree frogs (Anura: Rhacophoridae), with 387 species, display a remarkable diversity of reproductive modes – aquatic breeding, terrestrial gel nesting, terrestrial foam nesting and terrestrial direct development. The evolution of these modes has until now remained poorly studied in the context of recent phylogenies for the clade. Here, we use newly obtained DNA sequences from three nuclear and two mitochondrial gene fragments, together with previously published sequence data, to generate a well‐resolved phylogeny from which we determine major patterns of reproductive‐mode evolution. We show that basal rhacophorids have fully aquatic eggs and larvae. Bayesian ancestral‐state reconstructions suggest that terrestrial gel‐encapsulated eggs, with early stages of larval development completed within the egg outside of water, are an intermediate stage in the evolution of terrestrial direct development and foam nesting. The ancestral forms of almost all currently recognized genera (except the fully aquatic basal forms) have a high likelihood of being terrestrial gel nesters. Direct development and foam nesting each appear to have evolved at least twice within Rhacophoridae, suggesting that reproductive modes are labile and may arise multiple times independently. Evolution from a fully aquatic reproductive mode to more terrestrial modes (direct development and foam nesting) occurs through intermediate gel nesting ancestral forms. This suggests that gel nesting is not only a possible transitional state for the evolution of terrestriality, but also that it is a versatile reproductive mode that may give rise to other terrestrial reproductive modes. Evolution of foam nesting may have enabled rhacophorids to lay a larger number of eggs in more open and drier habitats, where protection from desiccation is important. Terrestrial direct development allows frogs to lay eggs independent of bodies of water, in a diversity of humid habitats, and may represent a key innovation that facilitated the evolution of nearly half of all known rhacophorid species.