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The evolution of life cycles forms the subject of numerous studies on extant organisms, but is rarely documented in the fossil record. Here, I analyze patterns of development in time-averaged samples of late Carboniferous and early Permian amphibians, and compare them to paleoecological patterns derived from the same deposits located within a large sedimentary basin (Saar-Nahe, Germany). In 300-297 million years (myr) old Sclerocephalus haeuseri (1-1.7 m), adult size, morphology, and the course of ontogeny varied with respect to the habitats in which the species existed. These differences are best exemplified by ontogenetic trajectories, which reveal a full range of modifications correlating with environmental parameters (lake properties, food resources, competitors). In a 2- to 3-myr-long interval, six different lake habitats were inhabited by this species, which responded to changes by modification of growth rate, adult size, developmental sequence, skeletal features, prey preference, and relative degree of terrestriality.

Among the diverse clade of Paleozoic dissorophoid amphibians, the small, terrestrial amphibamids and the neotenic branchiosaurids have frequently been suggested as possible antecedents of either all or some of the modern amphibian clades. Classically, amphibamids and branchiosaurids have been considered to represent distinct, but closely related clades within dissorophoids, but despite their importance for the controversial lissamphibian origins, a comprehensive phylogenetic analysis of small dissorophoids has thus far not been attempted. On the basis of an integrated data set, the relationships of amphibamids and branchiosaurids were analyzed using parsimony and Bayesian approaches. Both groups represent miniaturized forms and it was tested whether similar developmental pathways, associated with miniaturization, lead to an artificial close relationship of branchiosaurids and amphibamids. Moreover, the fit of the resulting tree topologies to the distribution of fossil taxa in the stratigraphic rock record was assessed as an additional source of information. The results show that characters associated with a miniaturized morphology are not responsible for the close clustering of branchiosaurids and amphibamids. Instead, all analyses invariably demonstrate a monophyletic clade of branchiosaurids highly nested within derived amphibamids, indicating that branchiosaurids represent a group of secondarily neotenic amphibamid dissorophoids. This understanding of the phylogenetic relationships of small dissorophoid amphibians provides a new framework for the discussion of their evolutionary history and the evolution of characters shared by branchiosaurids and/or amphibamids with modern amphibian taxa.

The Branchiosauridae was a clade of small amphibians from the Permo-Carboniferous with an overall salamander-like appearance. The clade is distinguished by an extraordinary fossil record that comprises hundreds of well-preserved specimens, representing a wide range of ontogenetic stages. Branchiosaurids had external gills and weakly ossified skeletons, and due to this larval appearance their status as neotenic (perennibranchiate) forms has long been accepted. Despite their extensive fossil record large specimens with an adult morphology appeared to be lacking altogether, but recently two adult specimens were identified in a rich sample of Apateon gracilis collected in the 19th century from a locality near Dresden, Saxony. These specimens are unique among branchiosaurids in showing a high level of ossification, including bones that have never been reported in a branchiosaur. These highlight the successive formation of features believed to indicate terrestrial locomotion, as well as feeding on larger prey items. Moreover, these transformations occurred in a small time window (whereas the degree of size increase is used as a proxy of time) and the degree of concentration of developmental events in branchiosaurids is unique among tetrapods outside the lissamphibians. These specimens are compared with large adults of the neotenic branchiosaurid Apateon caducus from the Saar-Nahe Basin, which despite their larger body size lack the features found in the adult A. gracilis specimens. These specimens give new insight into patterns of metamorphosis (morphological transformation) in branchiosaurids that are believed to be correlated to a change of habitat, and clearly show that different life-history pathways comparable to those of modern salamanders were already established in this Paleozoic clade.