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Despite the wide range of shapes and sizes that accompany a vast variety of functions, the development of tetrapod limbs follows a conservative pattern of de novo condensation, branching, and segmentation. Development of the zeugopodium and digital arch typically occurs in a posterior to anterior sequence, referred to as postaxial dominance, with a digital sequence of 4-3-5-2-1. The only exception to this pattern in all of living Tetrapoda can be found in salamanders, which display a preaxial dominance in limb development, a de novo condensation of a basale commune (distal carpal/tarsal 1+2) and a precoccial development of digits I and II. These divergent patterns have puzzled researchers for over a century leading to various explanatory hypotheses. Despite many advances in research on tetrapod limb development, the divergent evolution of these two pathways and its causes are still not understood. Based on an extensive ontogenetic series we investigated the pattern of limb development of the 300 Ma old branchiosaurid amphibian Apateon. This revealed a preaxial dominance in limb development that was previously believed to be unique and derived for modern salamanders. The Branchiosauridae are favored as close relatives of extant salamanders in most phylogenetic hypotheses of the highly controversial origins and relationships of extant amphibians. The findings provide new insights into the evolution of developmental pathways in tetrapod limb development, the relationships of modern amphibians with possible Paleozoic antecedents, and their initial timing of divergence.

In vertebrates, the ontogeny of the bony skull forms a particularly complex part of embryonic development. Although this area used to be restricted to neontology, recent discoveries of fossil ontogenies provide an additional source of data. One of the most detailed ossification sequences is known from Permo-Carboniferous amphibians, the branchiosaurids. These temnospondyls form a near-perfect link between the piscine osteichthyans and the various clades of extant tetrapods, retaining a full complement of dermal bones in the skull. For the first time, the broader evolutionary significance of these event sequences is analyzed, focusing on the identification of sequence heterochronies. A set of 120 event pairs was analyzed by event pair cracking, which helped identify active movers. A cladistic analysis of the event pair data was also carried out, highlighting some shared patterns between widely divergent clades of tetrapods. The analyses revealed an unexpected degree of similarity between the widely divergent taxa. Most interesting is the apparently modular composition of the cranial sequence: five clusters of bones were discovered in each of which the elements form in the same time window: (1) jaw bones, (2) marginal palatal elements, (3) circumorbital bones, (4) skull roof elements, and (5) neurocranial ossifications. In the studied taxa, these "modules" have in most cases been shifted fore and back on the trajectory relative to the Amia sequence, but did not disintegrate. Such "modules" might indicate a high degree of evolutionary limitation (constraint).

The phylogenetic positions of frogs, salamanders, and caecilians have been difficult to establish. Data matrices based primarily on Paleozoic taxa support a monophyletic origin of all Lissamphibia but have resulted in widely divergent hypotheses of the nature of their common ancestor. Analysis that concentrates on the character states of the stem taxa of the extant orders, in contrast, suggests a polyphyletic origin from divergent Paleozoic clades. Comparison of patterns of larval development in Paleozoic and modern amphibians provides a means to test previous phylogenies based primarily on adult characteristics. This proves to be highly informative in the case of the origin of salamanders. Putative ancestors of salamanders are recognized from the Permo-Carboniferous boundary of Germany on the basis of ontogenetic changes observed in fossil remains of larval growth series. The entire developmental sequence from hatching to metamorphosis is revealed in an assemblage of over 600 specimens from a single locality, all belonging to the genus Apateon. Apateon forms the most speciose genus of the neotenic temnospondyl family Branchiosauridae. The sequence of ossification of individual bones and the changing configuration of the skull closely parallel those observed in the development of primitive living salamanders. These fossils provide a model of how derived features of the salamander skull may have evolved in the context of feeding specializations that appeared in early larval stages of members of the Branchiosauridae. Larvae of Apateon share many unique derived characters with salamanders of the families Hynobiidae, Salamandridae, and Ambystomatidae, which have not been recognized in any other group of Paleozoic amphibians.