The origin of the limbless caecilians remains a lasting question in vertebrate evolution. Molecular phylogenies and morphology support that caecilians are the sister taxon of batrachians (frogs and salamanders), from which they diverged no later than the early Permian. Although recent efforts have discovered new, early members of the batrachian lineage, the record of pre-Cretaceous caecilians is limited to a single species, The position of within tetrapod phylogeny is controversial, as it already acquired the specialized morphology that characterizes modern caecilians by the Jurassic. Here, we report on a small amphibian from the Upper Triassic of Colorado, United States, with a mélange of caecilian synapomorphies and general lissamphibian plesiomorphies. We evaluated its relationships by designing an inclusive phylogenetic analysis that broadly incorporates definitive members of the modern lissamphibian orders and a diversity of extinct temnospondyl amphibians, including stereospondyls. Our results place the taxon confidently within lissamphibians but demonstrate that the diversity of Permian and Triassic stereospondyls also falls within this group. This hypothesis of caecilian origins closes a substantial morphologic and temporal gap and explains the appeal of morphology-based polyphyly hypotheses for the origins of Lissamphibia while reconciling molecular support for the group's monophyly. Stem caecilian morphology reveals a previously unrecognized stepwise acquisition of typical caecilian cranial apomorphies during the Triassic. A major implication is that many Paleozoic total group lissamphibians (i.e., higher temnospondyls, including the stereospondyl subclade) fall within crown Lissamphibia, which must have originated before 315 million years ago.

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In addition to their devastating effects on global biodiversity, mass extinctions have had a long-term influence on the history of life by eliminating dominant lineages that suppressed ecological change. Here, we test whether the end-Permian mass extinction (252.3 Ma) affected the distribution of tetrapod faunas within the southern hemisphere and apply quantitative methods to analyze four components of biogeographic structure: connectedness, clustering, range size, and endemism. For all four components, we detected increased provincialism between our Permian and Triassic datasets. In southern Pangea, a more homogeneous and broadly distributed fauna in the Late Permian (Wuchiapingian, ∼257 Ma) was replaced by a provincial and biogeographically fragmented fauna by Middle Triassic times (Anisian, ∼242 Ma). Importantly in the Triassic, lower latitude basins in Tanzania and Zambia included dinosaur predecessors and other archosaurs unknown elsewhere. The recognition of heterogeneous tetrapod communities in the Triassic implies that the end-Permian mass extinction afforded ecologically marginalized lineages the ecospace to diversify, and that biotic controls (i.e., evolutionary incumbency) were fundamentally reset. Archosaurs, which began diversifying in the Early Triassic, were likely beneficiaries of this ecological release and remained dominant for much of the later Mesozoic.biogeography | complex networks | macroevolution | biotic recovery | paleoecology M ass extinctions are thought to reshape the composition and ecological structure of communities on a scale unparalleled by background extinction (1, 2). Within the terrestrial realm, the replacement of dinosaur-dominated communities by those of mammals after the end-Cretaceous extinction is perhaps the best-known example of such wholesale faunal reshuffling. By contrast, understanding the effects of the more massive endPermian extinction on terrestrial community structure has been hampered by the paucity of high-quality geographic data, with nearly all studies restricted to sequences from single basins in Russia (3, 4) or South Africa (5-8). Although some broad similarities have emerged, such as the progressive aridification of the two basins (3, 8) and a heightened diversity of temnospondyl amphibians in the recovery interval (9), limited geographic sampling in each hemisphere constrains the ability of paleontologists to distinguish regional patterns from those characteristic of the individual basins. Moreover, these broadly separated areas are uninformative regarding large-scale (i.e., continentlevel) patterns of faunal evolution.Here we examine tetrapod faunal composition in five fossiliferous areas in southern Pangea approximately 5 million years before and 10 million years after the end-Permian mass extinction. We analyze changes in biogeographic structure with four metrics of taxon occurrence data. First, biogeographic connectedness (BC) quantifies the proportion of taxon-locality occurrences relative to the maximum number of such occurrences possible. Thus...

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