A Triassic stem-salamander from Kyrgyzstan and the origin of salamanders.

The origin of extant amphibians remains largely obscure, with only a few early Mesozoic stem taxa known, as opposed to a much better fossil record from the mid-Jurassic on. In recent time, anurans have been traced back to Early Triassic forms and caecilians have been traced back to the Late Jurassic Eocaecilia, both of which exemplify the stepwise acquisition of apomorphies. Yet the most ancient stem-salamanders, known from mid-Jurassic rocks, shed little light on the origin of the clade. The gap between salamanders and other lissamphibians, as well as Paleozoic tetrapods, remains considerable. Here we report a new specimen of Triassurus sixtelae, a hitherto enigmatic tetrapod from the Middle/Late Triassic of Kyrgyzstan, which we identify as the geologically oldest stem-group salamander. This sheds light not only on the early evolution of the salamander body plan, but also on the origin of the group as a whole. The new, second specimen is derived from the same beds as the holotype, the Madygen Formation of southwestern Kyrgyzstan. It reveals a range of salamander characters in this taxon, pushing back the rock record of urodeles by at least 60 to 74 Ma (Carnian-Bathonian). In addition, this stem-salamander shares plesiomorphic characters with temnospondyls, especially branchiosaurids and amphibamiforms.

A Middle Triassic stem-turtle and the evolution of the turtle body plan.

The origin and early evolution of turtles have long been major contentious issues in vertebrate zoology. This is due to conflicting character evidence from molecules and morphology and a lack of transitional fossils from the critical time interval. The ∼220-million-year-old stem-turtle Odontochelys from China has a partly formed shell and many turtle-like features in its postcranial skeleton. Unlike the 214-million-year-old Proganochelys from Germany and Thailand, it retains marginal teeth and lacks a carapace. Odontochelys is separated by a large temporal gap from the ∼260-million-year-old Eunotosaurus from South Africa, which has been hypothesized as the earliest stem-turtle. Here we report a new reptile, Pappochelys, that is structurally and chronologically intermediate between Eunotosaurus and Odontochelys and dates from the Middle Triassic period (∼240 million years ago). The three taxa share anteroposteriorly broad trunk ribs that are T-shaped in cross-section and bear sculpturing, elongate dorsal vertebrae, and modified limb girdles. Pappochelys closely resembles Odontochelys in various features of the limb girdles. Unlike Odontochelys, it has a cuirass of robust paired gastralia in place of a plastron. Pappochelys provides new evidence that the plastron partly formed through serial fusion of gastralia. Its skull has small upper and ventrally open lower temporal fenestrae, supporting the hypothesis of diapsid affinities of turtles.

Cranial shape evolution of extant and fossil crocodile newts and its relation to reproduction and ecology.

The diversity of the vertebrate cranial shape of phylogenetically related taxa allows conclusions on ecology and life history. As pleurodeline newts (the genera Echinotriton, Pleurodeles and Tylototriton) have polymorphic reproductive modes, they are highly suitable for following cranial shape evolution in relation to reproduction and environment. We investigated interspecific differences externally and differences in the cranial shape of pleurodeline newts via two-dimensional geometric morphometrics. Our analyses also included the closely related but extinct genus Chelotriton to better follow the evolutionary history of cranial shape. Pleurodeles was morphologically distinct in relation to other phylogenetically basal salamanders. The subgenera within Tylototriton (Tylototriton and Yaotriton) were well separated in morphospace, whereas Echinotriton resembled the subgenus Yaotriton more than Tylototriton. Oviposition site choice correlated with phylogeny and morphology. Only the mating mode, with a random distribution along the phylogenetic tree, separated crocodile newts into two morphologically distinct groups. Extinct Chelotriton likely represented several species and were morphologically and ecologically more similar to Echinotriton and Yaotriton than to Tylototriton subgenera. Our data also provide the first comprehensive morphological support for the molecular phylogeny of pleurodeline newts.

Amphibian skull evolution: The developmental and functional context of simplification, bone loss and heterotopy.

Despite their divergent morphology, extant and extinct amphibians share numerous features in the timing and spatial patterning of dermal skull elements. Here, I show how the study of these features leads to a deeper understanding of morphological evolution. Batrachians (salamanders and frogs) have simplified skulls, with dermal bones appearing rudimentary compared with fossil tetrapods, and open cheeks resulting from the absence of other bones. The batrachian skull bones may be derived from those of temnospondyls by truncation of the developmental trajectory. The squamosal, quadratojugal, parietal, prefrontal, parasphenoid, palatine, and pterygoid form rudimentary versions of their homologs in temnospondyls. In addition, failure to ossify and early fusion of bone primordia both result in the absence of further bones that were consistently present in Paleozoic tetrapods. Here, I propose a new hypothesis explaining the observed patterns of bone loss and emargination in a functional context. The starting observation is that jaw-closing muscles are arranged in a different way than in ancestors from the earliest ontogenetic stage onwards, with muscles attaching to the dorsal side of the frontal, parietal, and squamosal. The postparietal and supratemporal start to ossify in a similar way as in branchiosaurids, but are fused to neighboring elements to form continuous attachment areas for the internal adductor. The postfrontal, postorbital, and jugal fail to ossify, as their position is inconsistent with the novel arrangement of adductor muscles. Thus, rearrangement of adductors forms the common theme behind cranial simplification, driven by an evolutionary flattening of the skull in the batrachian stem. J. Exp. Zool. (Mol. Dev. Evol.) 9999B: XX-XX, 2014. © 2014 Wiley Periodicals, Inc.

Amphibian skull evolution: the developmental and functional context of simplification, bone loss and heterotopy.

Despite their divergent morphology, extant and extinct amphibians share numerous features in the timing and spatial patterning of dermal skull elements. Here, I show how the study of these features leads to a deeper understanding of morphological evolution. Batrachians (salamanders and frogs) have simplified skulls, with dermal bones appearing rudimentary compared with fossil tetrapods, and open cheeks resulting from the absence of other bones. The batrachian skull bones may be derived from those of temnospondyls by truncation of the developmental trajectory. The squamosal, quadratojugal, parietal, prefrontal, parasphenoid, palatine, and pterygoid form rudimentary versions of their homologs in temnospondyls. In addition, failure to ossify and early fusion of bone primordia both result in the absence of further bones that were consistently present in Paleozoic tetrapods. Here, I propose a new hypothesis explaining the observed patterns of bone loss and emargination in a functional context. The starting observation is that jaw-closing muscles are arranged in a different way than in ancestors from the earliest ontogenetic stage onwards, with muscles attaching to the dorsal side of the frontal, parietal, and squamosal. The postparietal and supratemporal start to ossify in a similar way as in branchiosaurids, but are fused to neighboring elements to form continuous attachment areas for the internal adductor. The postfrontal, postorbital, and jugal fail to ossify, as their position is inconsistent with the novel arrangement of adductor muscles. Thus, rearrangement of adductors forms the common theme behind cranial simplification, driven by an evolutionary flattening of the skull in the batrachian stem.

The oldest known rhynchocephalian reptile from the Middle Triassic (Ladinian) of Germany and its phylogenetic position among Lepidosauromorpha.

Skeletal remains of a small lepidosaurian reptile from the Middle Triassic (Ladinian: Longobardian) Erfurt Formation, exposed in a commercial limestone quarry near Vellberg (Germany), represent the oldest rhynchocephalian known to date. The new taxon, Wirtembergia hauboldae, is diagnosed by the following combination of features: Premaxilla with four teeth, first being largest and decreasing in size from first to fourth. Jugal with tiny, spur-like posterior process. Lateral surface of dentary strongly convex dorsoventrally for much of length of bone, bearing distinct longitudinal ridge and sculpturing in large specimens. Coronoid eminence of dentary low, subrectangular, and with dorsoventrally concave lateral surface in larger specimens. Dentition with pleurodont anterior and acrodont posterior teeth. Posterior (=additional) teeth with (in side view) triangular, at mid-crown level labiolingually somewhat flattened crowns, and with oval bases. Phylogenetic analysis recovered the new rhynchocephalian as the earliest-diverging member of its clade known to date.

The evolution of larvae in temnospondyls and the stepwise origin of amphibian metamorphosis.

The question of what the ancient life cycle of tetrapods was like forms a key component in understanding the origin of land vertebrates. The existence of distinct larval forms, as exemplified by many lissamphibians, and their transformation into adults is an important aspect in this field. The temnospondyls, the largest clade of Palaeozoic-Mesozoic non-amniote tetrapods, covered a wide ecomorphological range from fully aquatic to terrestrial taxa. In various species, rich ontogenetic data have accumulated over the past 130 years, permitting the study of early phases of temnospondyl development. In temnospondyls, eight ontogenetic phases have been identified in which the skeleton formed. In branchiosaurids and the eryopiform Sclerocephalus, large parts of the ossification sequence are now known. Most taxa in which small specimens are preserved had aquatic larvae with external gills that superficially resemble larval salamanders. In the edopoids, dvinosaurs, and eryopiforms, the larvae developed slowly, with incompletely ossified axial and appendicular skeletons, but possessed a fast-developing dermal skull with strong teeth. Irrespective of adult terrestriality or a fully aquatic life, there was no drastic transformation during later ontogeny, but a slow and steady acquisition of adult features. In dissorophoids, the limbs developed at a much faster pace, whereas skull formation was slowed down, especially in the amphibamiforms, and culminating in the neotenic Branchiosauridae. In the zatracheid Acanthostomatops, slow but profound transformation led to a fully terrestrial adult. The basal dissorophoid Stegops retained rapid development of dermal skull bones and established a fully dentigerous, strongly ossified palate early. In Micromelerpeton, formation of the last skull bones was slightly delayed and metamorphosis remained a long and steady phase of morphological transformations. In amphibamiforms, metamorphosis became more drastic, with an increasing number of events packed into a short phase of ontogeny. This is exemplified by Apateon, Platyrhinops, and Amphibamus in which this condensation was maximised. We distinguish three different types of metamorphosis (morphological, ecological and drastic) that evolved cumulatively in early tetrapods and within temnospondyls.

Paleobiological implications of chevron pathology in the sauropodomorph Plateosaurus trossingensis from the Upper Triassic of SW Germany.

Paleopathology, the study of diseases and injuries from the fossil record, allows for a unique view into the life of prehistoric animals. Pathologies have nowadays been described in nearly all groups of fossil vertebrates, especially dinosaurs. Despite the large number of skeletons, pathologies had never been reported in the sauropodomorph Plateosaurus trossingensis. Here we describe the first pathologies of Plateosaurus using two individuals with pathologies in the chevrons of the tail, from the Upper Triassic of Trossingen, SW Germany. The two specimens each contain three consecutive pathological chevrons. Our results show that the pathologies were caused by external trauma in one individual and potentially tendinous trauma in the other. Healing of the lesions allowed survival of both animals. Using additional pathological specimens found in other collections and from multiple localities, we observe that 14.8% of all individuals of Plateosaurus contain pathologies within their chevrons, suggesting it was a vulnerable bone.

The girdles of the oldest fossil turtle, Proterochersis robusta, and the age of the turtle crown.

BACKGROUND: Proterochersis robusta from the Late Triassic (Middle Norian) of Germany is the oldest known fossil turtle (i.e. amniote with a fully formed turtle shell), but little is known about its anatomy. A newly prepared, historic specimen provides novel insights into the morphology of the girdles and vertebral column of this taxon and the opportunity to reassess its phylogenetic position. RESULTS: The anatomy of the pectoral girdle of P. robusta is similar to that of other primitive turtles, including the Late Triassic (Carnian) Proganochelys quenstedti, in having a vertically oriented scapula, a large coracoid foramen, a short acromion process, and bony ridges that connect the acromion process with the dorsal process, glenoid, and coracoid, and by being able to rotate along a vertical axis. The pelvic elements are expanded distally and suturally attached to the shell, but in contrast to modern pleurodiran turtles the pelvis is associated with the sacral ribs. CONCLUSIONS: The primary homology of the character "sutured pelvis" is unproblematic between P. robusta and extant pleurodires. However, integration of all new observations into the most complete phylogenetic analysis that support the pleurodiran nature of P. robusta reveals that this taxon is more parsimoniously placed along the phylogenetic stem of crown Testudines. All current phylogenetic hypotheses therefore support the basal placement of this taxon, imply that the sutured pelvis of this taxon developed independently from that of pleurodires, and conclude that the age of the turtle crown is Middle Jurassic.

Integration of molecules and new fossils supports a Triassic origin for Lepidosauria (lizards, snakes, and tuatara).

BACKGROUND: Lepidosauria (lizards, snakes, tuatara) is a globally distributed and ecologically important group of over 9,000 reptile species. The earliest fossil records are currently restricted to the Late Triassic and often dated to 227 million years ago (Mya). As these early records include taxa that are relatively derived in their morphology (e.g. Brachyrhinodon), an earlier unknown history of Lepidosauria is implied. However, molecular age estimates for Lepidosauria have been problematic; dates for the most recent common ancestor of all lepidosaurs range between approximately 226 and 289 Mya whereas estimates for crown-group Squamata (lizards and snakes) vary more dramatically: 179 to 294 Mya. This uncertainty restricts inferences regarding the patterns of diversification and evolution of Lepidosauria as a whole. RESULTS: Here we report on a rhynchocephalian fossil from the Middle Triassic of Germany (Vellberg) that represents the oldest known record of a lepidosaur from anywhere in the world. Reliably dated to 238-240 Mya, this material is about 12 million years older than previously known lepidosaur records and is older than some but not all molecular clock estimates for the origin of lepidosaurs. Using RAG1 sequence data from 76 extant taxa and the new fossil specimens two of several calibrations, we estimate that the most recent common ancestor of Lepidosauria lived at least 242 Mya (238-249.5), and crown-group Squamata originated around 193 Mya (176-213). CONCLUSION: A Early/Middle Triassic date for the origin of Lepidosauria disagrees with previous estimates deep within the Permian and suggests the group evolved as part of the faunal recovery after the end-Permain mass extinction as the climate became more humid. Our origin time for crown-group Squamata coincides with shifts towards warmer climates and dramatic changes in fauna and flora. Most major subclades within Squamata originated in the Cretaceous postdating major continental fragmentation. The Vellberg fossil locality is expected to become an important resource for providing a more balanced picture of the Triassic and for bridging gaps in the fossil record of several other major vertebrate groups.

A new Middle Triassic (Ladinian) trilophosaurid stem-archosaur from Germany increases diversity and temporal range of this clade.

We report the first trilophosaurid stem-archosaur from Central Europe, Rutiotomodon tytthos gen. et sp. nov., from the Middle Triassic (Ladinian) Erfurt Formation of Baden-Württemberg (Germany). It is currently known from two jaw fragments with distinctive teeth. The labiolingually wide but mesiodistally narrow maxillary and dentary teeth each have a large labial cusp from which an occlusal ridge extends lingually to a small lingual cusp. A mesial and a distal cingulum extend between the labial and lingual cusps. The mesial and distal faces of the labial cusp each bear three prominent, lingually curved apicobasal ridges (arrises). A referred partial dentary has an edentulous, expanded symphysis similar to the mandibular 'beak' in Trilophosaurus buettneri. A review of Coelodontognathus ricovi, from the Lower Triassic (Olenekian) of southwestern Russia, supports its referral to Trilophosauridae rather than Procolophonidae. Based on this reassessment and the new material from the Middle Triassic, the temporal range of trilophosaurids now spans nearly the entire Triassic Period, from the Olenekian to the Rhaetian. Trilophosaurids present craniodental features that indicate omnivory or herbivory with limited oral food processing. They were more diverse in terms of dental structure (and presumably diet) than previously assumed.

Using salamanders as model taxa to understand vertebrate feeding constraints during the late Devonian water-to-land transition.

The vertebrate water-to-land transition and the rise of tetrapods brought about fundamental changes for the groups undergoing these evolutionary changes (i.e. stem and early tetrapods). These groups were forced to adapt to new conditions, including the distinct physical properties of water and air, requiring fundamental changes in anatomy. Nutrition (or feeding) was one of the prime physiological processes these vertebrates had to successfully adjust to change from aquatic to terrestrial life. The basal gnathostome feeding mode involves either jaw prehension or using water flows to aid in ingestion, transportation and food orientation. Meanwhile, processing was limited primarily to simple chewing bites. However, given their comparatively massive and relatively inflexible hyobranchial system (compared to the more muscular tongue of many tetrapods), it remains fraught with speculation how stem and early tetrapods managed to feed in both media. Here, we explore ontogenetic water-to-land transitions of salamanders as functional analogues to model potential changes in the feeding behaviour of stem and early tetrapods. Our data suggest two scenarios for terrestrial feeding in stem and early tetrapods as well as the presence of complex chewing behaviours, including excursions of the jaw in more than one dimension during early developmental stages. Our results demonstrate that terrestrial feeding may have been possible before flexible tongues evolved. This article is part of the theme issue 'Food processing and nutritional assimilation in animals'.

Amphibian development in the fossil record.

Ontogenetic series of extinct taxa are extremely rare and when preserved often incomplete and difficult to interpret. However, the fossil record of amphibians includes a number of well-preserved ontogenetic sequences for temnospondyl and lepospondyl taxa, which have provided valuable information about the development of these extinct groups. Here we summarize the current knowledge on fossil ontogenies of amphibians, their potential and limitations for relationship assessments, and discuss the insights they have provided for our understanding of the anatomy, life history, and ecology of extinct amphibians.

Riedl's burden and the body plan: selection, constraint, and deep time.

Rupert Riedl's concept of burden forms a causal hypothesis on organismic integration and evolutionary constraints. Defined as the hierarchically nested interdependence of characters within the organism, burden was seen as (1) defining and conserving body plans and (2) constraining and directing evolutionary trajectories. A review of the components of the burden concept reveals important consistencies with the modern tenets of evo-devo. This concept differs from the current consensus of evolutionary theory in that it (1) grants evolution less options for changing tightly integrated, "locked-in" characters and (2) in deducing from this an ever decreasing freedom for evolution, with cyclism and typostrophism as resulting macroevolutionary phenomena. Despite these differences, I show that the burden concept was consistent with most major tenets of the Modern Synthesis, and Riedl attempted to explain patterns of large-scale evolutionary trends exclusively by microevolutionary (gradualistic) processes. The burden concept is fruitful and unique in its focus on hierarchically nested constraints and resembles the hierarchical architecture of gene regulatory networks. However, such networks are more high-dimensional and most of their components appear to be easier to evolve than Riedl's burden. Yet in combination with evolvability, a modified concept of burden might contribute substantially to the understanding of organismic integration and the long-term evolution of body plans.

Testing the impact of miniaturization on phylogeny: Paleozoic dissorophoid amphibians.

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.

A tiny new Middle Triassic stem-lepidosauromorph from Germany: implications for the early evolution of lepidosauromorphs and the Vellberg fauna.

The Middle Triassic was a time of major changes in tetrapod faunas worldwide, but the fossil record for this interval is largely obscure for terrestrial faunas. This poses a severe limitation to our understanding on the earliest stages of diversification of lineages representing some of the most diverse faunas in the world today, such as lepidosauromorphs (e.g., lizards and tuataras). Here, we report a tiny new lepidosauromorph from the Middle Triassic from Vellberg (Germany), which combines a mosaic of features from both early evolving squamates and rhynchocephalians, such as the simultaneous occurrence of a splenial bone and partial development of acrodonty. Phylogenetic analyses applying different optimality criteria, and combined morphological and molecular data, consistently recover the new taxon as a stem-lepidosauromorph, implying stem-lepidosauromorph species coinhabited areas comprising today's central Europe at the same time as the earliest known rhynchocephalians and squamates. It further demonstrates a more complex evolutionary scenario for dental evolution in early lepidosauromorphs, with independent acquisitions of acrodonty early in their evolutionary history. The small size of most terrestrial vertebrates from Vellberg is conspicuous, contrasting to younger Triassic deposits worldwide, but comparable to Early Triassic faunas, suggesting a potential long-lasting Lilliput effect in this fauna.

The armoured dissorophid Cacops from the Early Permian of Oklahoma and the exploitation of the terrestrial realm by amphibians.

Cacops, one of the most distinctive Paleozoic amphibians, is part of a clade of dissorophoid temnospondyls that diversified in the equatorial region of Pangea during the Late Carboniferous and Early Permian, persisting into the Late Permian in Central Russia and China. Dissorophids were a successful group of fully terrestrial, often spectacularly armoured predators, the only amphibians apparently able to coexist with amniotes when the latter started to dominate terrestrial ecosystems. In this paper, we describe excellent new skulls from the Early Permian of Oklahoma attributed to Cacops, Cacops morrisi sp. nov. and provide for the first time detailed information about this iconic dissorophid. These specimens show anatomical and ontogenetic features that will impact on future studies on the evolution of terrestriality in tetrapods. For example, the large, posteriorly closed tympanic embayment has fine striations on an otherwise smooth surface, documenting the oldest known clear evidence for the presence of a tympanic membrane in the fossil record, a structure that is used for hearing airborne sound in extant tetrapods. The skull of C. morrisi also has several features associated with predatory behaviour, indicating that this dissorophid may have been one of the top terrestrial predators of its time.

Microanatomy of the stem-turtle Pappochelys rosinae indicates a predominantly fossorial mode of life and clarifies early steps in the evolution of the shell.

Unlike any other tetrapod, turtles form their dorsal bony shell (carapace) not from osteoderms, but by contribution of the ribs and vertebrae that expand into the dermis to form plate-like shell components. Although this was known from embryological studies in extant turtles, important steps in this evolutionary sequence have recently been highlighted by the Triassic taxa Pappochelys, Eorhynchochelys and Odontochelys, and the Permian Eunotosaurus. The discovery of Pappochelys shed light on the origin of the ventral bony shell (plastron), which formed from enlarged gastralia. A major question is whether the turtle shell evolved in the context of a terrestrial or aquatic environment. Whereas Odontochelys was controversially interpreted as aquatic, a terrestrial origin of turtles was proposed based on evidence of fossorial adaptations in Eunotosaurus. We report palaeohistological data for Pappochelys, a taxon that exemplifies earlier evolutionary stages in the formation of the bony shell than Odontochelys. Bone histological evidence reveals (1) evolutionary changes in bone microstructure in ribs and gastralia approaching the turtle condition and (2) evidence for a predominantly amphibious or fossorial mode of life in Pappochelys, which support the hypothesis that crucial steps in the evolution of the shell occurred in a terrestrial rather than fully aquatic environment.

A relict basal tetrapod from Germany: first evidence of a Triassic chroniosuchian outside Russia.

Chroniosuchians are basal tetrapods nesting within the paraphyletic anthracosaurs and were so far only well known from the Permian and Triassic of Russia. In this study, we present evidence for their existence in the upper Middle Triassic of Germany, based on diagnostic osteoderms and vertebrae from the Kupferzell and Vellberg localities in southern Germany. The finds are most similar to Synesuchus, a Middle Triassic bystrowianid chroniosuchian from the Northern Ural Pechora region. They demonstrate that by Middle Triassic time, chroniosuchians were much more widespread than previously thought.

Metamorphosis and neoteny: alternative pathways in an extinct amphibian clade.

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.