Rainer R. Schoch

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

BackgroundLepidosauria (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.ResultsHere 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).ConclusionA 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.

Postcranial Anatomy of the Rauisuchian Archosaur Batrachotomus kupferzellensis

ABSTRACT Batrachotomus kupferzellensis is an upper Middle Triassic (Late Ladinian) rauisuchian archosaur. The postcranial skeleton of this species is well-represented by fossil material, including the holotype, from the localities of Kupferzell, Crailsheim and Vellberg-Eschenau in southern Germany, and is described here in detail for the first time. All postcranial elements are known except the interclavicle and parts of the carpus, manus, tarsus, pes and some osteoderm and axial elements. B. kupferzellensis is now one of the best-known rauisuchians and will be important in advancing understanding of the groups biology. A period of new anatomical and taxonomic work since 2000 has improved understanding of rauisuchians. Renewed effort in rauisuchian phylogenetics will benefit from these new data, but will also require a careful and detailed approach to character formulation.

The Sail-Backed Reptile Ctenosauriscus from the Latest Early Triassic of Germany and the Timing and Biogeography of the Early Archosaur Radiation

Background Archosaurs (birds, crocodilians and their extinct relatives including dinosaurs) dominated Mesozoic continental ecosystems from the Late Triassic onwards, and still form a major component of modern ecosystems (>10,000 species). The earliest diverse archosaur faunal assemblages are known from the Middle Triassic (c. 244 Ma), implying that the archosaur radiation began in the Early Triassic (252.3–247.2 Ma). Understanding of this radiation is currently limited by the poor early fossil record of the group in terms of skeletal remains. Methodology/Principal Findings We redescribe the anatomy and stratigraphic position of the type specimen of Ctenosauriscus koeneni (Huene), a sail-backed reptile from the Early Triassic (late Olenekian) Solling Formation of northern Germany that potentially represents the oldest known archosaur. We critically discuss previous biomechanical work on the ‘sail’ of Ctenosauriscus, which is formed by a series of elongated neural spines. In addition, we describe Ctenosauriscus-like postcranial material from the earliest Middle Triassic (early Anisian) Röt Formation of Waldhaus, southwestern Germany. Finally, we review the spatial and temporal distribution of the earliest archosaur fossils and their implications for understanding the dynamics of the archosaur radiation. Conclusions/Significance Comprehensive numerical phylogenetic analyses demonstrate that both Ctenosauriscus and the Waldhaus taxon are members of a monophyletic grouping of poposauroid archosaurs, Ctenosauriscidae, characterised by greatly elongated neural spines in the posterior cervical to anterior caudal vertebrae. The earliest archosaurs, including Ctenosauriscus, appear in the body fossil record just prior to the Olenekian/Anisian boundary (c. 248 Ma), less than 5 million years after the Permian–Triassic mass extinction. These earliest archosaur assemblages are dominated by ctenosauriscids, which were broadly distributed across northern Pangea and which appear to have been the first global radiation of archosaurs.

Skull ontogeny: developmental patterns of fishes conserved across major tetrapod clades

SUMMARY 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 Cranial Osteology and Feeding Ecology of the Metriorhynchid Crocodylomorph Genera Dakosaurus and Plesiosuchus from the Late Jurassic of Europe

Background Dakosaurus and Plesiosuchus are characteristic genera of aquatic, large-bodied, macrophagous metriorhynchid crocodylomorphs. Recent studies show that these genera were apex predators in marine ecosystems during the latter part of the Late Jurassic, with robust skulls and strong bite forces optimized for feeding on large prey. Methodology/Principal Findings Here we present comprehensive osteological descriptions and systematic revisions of the type species of both genera, and in doing so we resurrect the genus Plesiosuchus for the species Dakosaurus manselii. Both species are diagnosed with numerous autapomorphies. Dakosaurus maximus has premaxillary ‘lateral plates’; strongly ornamented maxillae; macroziphodont dentition; tightly fitting tooth-to-tooth occlusion; and extensive macrowear on the mesial and distal margins. Plesiosuchus manselii is distinct in having: non-amblygnathous rostrum; long mandibular symphysis; microziphodont teeth; tooth-crown apices that lack spalled surfaces or breaks; and no evidence for occlusal wear facets. Our phylogenetic analysis finds Dakosaurus maximus to be the sister taxon of the South American Dakosaurus andiniensis, and Plesiosuchus manselii in a polytomy at the base of Geosaurini (the subclade of macrophagous metriorhynchids that includes Dakosaurus, Geosaurus and Torvoneustes). Conclusions/Significance The sympatry of Dakosaurus and Plesiosuchus is curiously similar to North Atlantic killer whales, which have one larger ‘type’ that lacks tooth-crown breakage being sympatric with a smaller ‘type’ that has extensive crown breakage. Assuming this morphofunctional complex is indicative of diet, then Plesiosuchus would be a specialist feeding on other marine reptiles while Dakosaurus would be a generalist and possible suction-feeder. This hypothesis is supported by Plesiosuchus manselii having a very large optimum gape (gape at which multiple teeth come into contact with a prey-item), while Dakosaurus maximus possesses craniomandibular characteristics observed in extant suction-feeding odontocetes: shortened tooth-row, amblygnathous rostrum and a very short mandibular symphysis. We hypothesise that trophic specialisation enabled these two large-bodied species to coexist in the same ecosystem.

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.

THE AMPHIBAMID MICROPHOLIS FROM THE LYSTROSAURUS ASSEMBLAGE ZONE OF SOUTH AFRICA

Abstract The small temnospondyl Micropholis stowi from the Lower Triassic Lystrosaurus Assemblage Zone (Karoo Basin, South Africa) is redescribed on the basis of a larger sample of specimens, revealing a range of previously unknown anatomical features. Micropholis is recognized as a member of the Amphibamidae, representing both the last occurrence and the only Gondwanan member of this clade. (1) The largest specimens have an elongated narrow snout, larger otic notches, prominent quadrate processes, and wide cheeks giving the skull a marked lateral curvature. (2) Palpebral ossifications include a fabric of numerous polygonal ossicles contouring the size and position of the eyeballs. (3) The palate houses large fangs with inwardly curving crowns, an elaborate basicranial region resembling closely that of Amphibamus and Doleserpeton, and an extremely reduced rod-like palatine and ectopterygoid approaching the derived condition seen in other amphibamids. (4) In the postcranial skeleton, the greatly extended transverse processes, the morphology of the interclavicle and scapulocoracoid, and the structure of the pelvis and tail are characteristic features of Micropholis some of which are quite similar to the condition in Eoscopus and Platyrhinops. It is evident that two morphs of Micropholis, which differ in skull width and palatal dentition, are present in the Karoo Basin. Phylogenetic analysis suggests Micropholis to be nested within the amphibamid dissorophoids as the basal-most taxon. Within the Amphibamidae, Doleserpeton and Amphibamus are sister groups, nested with successive sister taxa Platyrhinops and Eoscopus. In particular, the miniaturized genera Doleserpeton and Amphibamus share a range of derived character-states with Micropholis, but the latter lacks the pedicellate dentition and has a more plesiomorphic posterior skull table and parasphenoid, suggesting that some of the shared derived states may have been acquired independently during the long isolated evolution of the Micropholis lineage in the Southern Hemisphere. In the present analysis, Micromelerpeton is the most basal offshoot of the dissorophoids, followed by a grade towards amphibamids formed by the following successive sister groups (from the base crownwards): (1) the trematopids, (2) Ecolsonia, and (3) the Dissorophidae, the latter being the sister taxon of all analyzed amphibamids.

METAMORPHOSIS AND NEOTENY: ALTERNATIVE PATHWAYS IN AN EXTINCT AMPHIBIAN CLADE

Abstract 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) froms 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 largetr 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 estabilshed in this Paleozoic clade.

The evolution of major temnospondyl clades: an inclusive phylogenetic analysis

Phylogenetic analysis of a large dataset (72 taxa, 212 characters) focuses on the in-group relationships of temnospondyls, the largest lower tetrapod clade. Representatives of all clades and grades were considered, spanning the entire stratigraphical range of temnospondyls from the Early Carboniferous through to the Early Cretaceous. Several major groups are defined phylogenetically (node or branch-based) rather than by apomorphies. The following groups were unequivocally found to be monophyletic: Edopoidea (node), Dvinosauria (stem, excl. Brachyopidae), Dissorophoidea (node), Eryopidae (stem), and Stereospondyli (node). The latter encompass three well-defined, branch-based taxa: Rhinesuchidae, Trematosauria and Capitosauria. Trematosauria (stem) contain Trematosauroidea (node), which includes the classic trematosaurids, metoposaurids, and possibly part of the rhytidosteids (Peltostega) but their in-group relationships remain unsettled; most other short-snouted stereospondyls (chigutisaurids, brachyopids, Laidleria and the plagiosaurids) are probably monophyletic and likely nest in some form with trematosauroids. Capitosauria (stem) include the Capitosauroidea (node) spanned by Parotosuchus and Mastodonsaurus, with the successive stem taxa Edingerella, Benthosuchus, Wetlugasaurus and Watsonisuchus. In all variant analyses, edopoids form the basalmost temnospondyl clade, followed by a potential clade (or grade) of small terrestrial taxa containing Balanerpeton and Dendrerpeton (‘Dendrerpetontidae’). All taxa higher than Edopoidea are suggested to form the monophyletic stem taxon Eutemnospondyli, tax. nov. The remainder of Temnospondyli fall into four robust and undisputed clades: (1) Dvinosauria; (2) Zatracheidae plus Dissorophoidea; (3) Eryopidae; and (4) Stereospondyli. These taxa are together referred to as Rhachitomi (node). Eryopidae and Stereospondylomorpha are probably monophyletic, here referred to as Eryopiformes (tax. nov.). The position of Dissorophoidea + Zatracheidae is still ambiguous; it may either form the sister taxon of Dvinosauria, or nest between Dvinosauria and Eryopiformes, whereas there is no support for Euskelia (Dissorophoidea + Eryopidae) after basal taxa of each clade are better understood. http://zoobank.org/urn:lsid:zoobank.org:pub:075B0FB3-8FF7-4247-BA49-424948F38EBE

Early larval ontogeny of the Permo‐Carboniferous temnospondyl Sclerocephalus

The early larval development of the temnospondyl Sclerocephalus sp. is analyzed, based on 38 specimens from the Lower Rotliegend (Permo-Carboniferous boundary) of the Saar-Nahe Basin (south-west Germany). The study focuses on the smallest larval specimens, which exemplify changes in both proportions and ossification patterns. In comparison with dissorophoid larvae, the skull ossifies more fully and at a much faster rate; the smallest specimens already have completely formed circumorbital bones that are sutured throughout. Sculpturing undergoes two marked changes, first from uniformly pitted to pits of variable size and regional differentiation, and finally to the origin of ridges. The palate of small larvae differs from that of larger specimens in patterns of dentition, having more teeth including a denticle field on the cultriform process. The mandible of small larvae is described for the first time, being narrower than in adults and having three dentigerous coronoid elements. The smallest specimens have poorly ossified neural arches, lack vertebral centra, and have faintly ossified humeri, femora, and very poorly developed distal elements. The posterior ribs, metapodia, and phalanges appeared after the dermal elements of the pectoral girdle, whereas the scapulocoracoid and ischium are absent throughout the larval period. Early growth and differentiation of the limbs and the ilium illustrates the developmental patterning of the appendages, which proceeded from proximal to distal. Dermal squamation is uniform in small stages, consisting of round or oval osteoderms with pronounced growth rings; in large larvae, they start to differentiate in certain body regions.