Temnospondyl ontogeny and phylogeny, a window into terrestrial ecosystems during the Permian-Triassic mass extinction.

摘要

Temnospondyls are the most species-rich group of early amphibians, but species-level phylogenetic analyses of this large clade have so far only incompletely sampled the group. This study represents the largest and most comprehensive species-level phylogenetic study of Temnospondyli, sampling 99 taxa for 297 morphological characters from all seven continents through nearly 170 million years of their evolutionary history. Results of this analysis support the monophyly of several clades. Phylogenetic definitions are updated and three new clades names are proposed: Eutemnospondyli, Neostereospondyli, and Latipalata. Major splits within temnospondyl evolution are recovered at the base of Eutemnospondyli (Euskelia and Limnarchia) and Neostereospondyli (Capitosauria and Trematosauria). Archegosauriodea is recovered within Euskelia. Dendrerpeton is recovered as the immediate sister taxon of Dissorophoidea, not Eryopoidea. This arrangement suggests that for subclade-level analyses of dissorophoids, which bear on the 'Temnospondyl Hypothesis' for a putative origin of Lissamphibia within dissorophoids, the convention of rooting on Dendrerpeton and including eyropoids in the ingroup should be re-evaluated in light of the new temnospondyl topology.;Study of the tempo and mode of evolution within temnospondyl amphibians has been limited in the past by the availability of a clade-wide, species-level phylogenetic analysis. The phylogenetic dataset generated by this study has allowed for investigation into rates of origination and extinction amongst this long-lived group at a scale not previously available for exploration. Extinction rate and origination rate, when calculated strictly from stratigraphic data, showed a high correlation with the number of sampled localities, indicating a strong influence on this evolutionary signal by sampling and rock record biases. But when rates were augmented with phylogenetic data, four periods of increased lineage origination are discernible from the Pennsylvanian to the Early Triassic. The largest of these origination events coincides with the Permo-Triassic mass extinction, indicating that amphibians were taking advantage of favorable conditions during the largest biological crisis in the Phanerozoic record.;Temnospondyl amphibians are the second most abundant fossil vertebrates in the Permo-Triassic Karoo Basin of South Africa. Paleohistological investigation of these amphibians was hampered by small sample size and taxa available for sampling. Incorporation of paleohistologic data from other analyses helped to alleviate this problem; however, Temnospondyli remains under sampled in paleohistological analyses. Results show cyclic growth and a lifespan of thirty years or more in basal stereospondyls, convergence to sustained, non-cyclic growth in terrestrial temnospondyls, support findings based on gross morphology that Lydekkerina is a terrestrial stereospondyl, and suggest that ribs are a viable source of skeletochronologic information in temnospondyls and should serve as preferred material when proximal limb diaphyses are not available. Sustained, azonal growth in Micropholis is unlike that of Apateon or extant caudatans, suggesting a possible adaptation to local conditions in the earliest Triassic of Gondwana.