A new sphenodontian (Diapsida: Lepidosauria) from the Upper Triassic (Norian) of Germany and its implications for the mode of sphenodontian evolution.

The Arnstadt Formation of Saxony-Anhalt, Germany has yielded some of Germany's most substantial finds of Late Triassic tetrapods, including the sauropodomorph Plateosaurus and the stem-turtle Proganochelys quenstedti. Here, we describe an almost complete skull of a new sphenodontian taxon from this formation (Norian, 227-208 Ma), making it the oldest known articulated sphenodontian skull from Europe and one of the oldest in the world. The material is represented by the dermal skull roof and by the complete maxilla and temporal region, as well as parts of the palate, braincase, and lower jaw. A phylogenetic assessment recovers it as a basal sphenodontian closely related to Planocephalosaurus robinsonae and to Eusphenodontia, making it the earliest-diverging sphenodontian known with an articulated skull. Its cranial anatomy is generally similar to the well-known Diphydontosaurus avonis from the Rhaetian of England, showing that this successful phenotype was already established in the clade around 10 myr earlier than assumed. An analysis of evolutionary change rates recovers high rates of evolution in basal sphenodontians, with decreasing rates throughout the evolution of the group. However, contrary to previous studies, reversals in this trend were identified, indicating additional peaks of evolutionary change. These results improve our understanding of the early sphenodontian diversity in Europe, providing critical information on evolutionary trends throughout the history of the clade and sparking renewed interest in its evolution.

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.

A new Carboniferous edaphosaurid and the origin of herbivory in mammal forerunners.

Herbivory evolved independently in several tetrapod lineages during the Late Carboniferous and became more widespread throughout the Permian Period, eventually leading to the basic structure of modern terrestrial ecosystems. Here we report a new taxon of edaphosaurid synapsid based on two fossils recovered from the Moscovian-age cannel coal of Linton, Ohio, which we interpret as an omnivore-low-fibre herbivore. Melanedaphodon hovaneci gen. et sp. nov. provides the earliest record of an edaphosaurid to date and is one of the oldest known synapsids. Using high-resolution X-ray micro-computed tomography, we provide a comprehensive description of the new taxon that reveals similarities between Late Carboniferous and early Permian (Cisuralian) members of Edaphosauridae. The presence of large bulbous, cusped, marginal teeth alongside a moderately-developed palatal battery, distinguishes Melanedaphodon from all other known species of Edaphosauridae and suggests adaptations for processing tough plant material already appeared among the earliest synapsids. Furthermore, we propose that durophagy may have provided an early pathway to exploit plant resources in terrestrial ecosystems.

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.

Arctic ice and the ecological rise of the dinosaurs.

Abundant lake ice-rafted debris in Late Triassic and earliest Jurassic strata of the Junggar Basin of northwestern China (paleolatitude ~71°N) indicates that freezing winter temperatures typified the forested Arctic, despite a persistence of extremely high levels of atmospheric Pco2 (partial pressure of CO2). Phylogenetic bracket analysis shows that non-avian dinosaurs were primitively insulated, enabling them to access rich deciduous and evergreen Arctic vegetation, even under freezing winter conditions. Transient but intense volcanic winters associated with massive eruptions and lowered light levels led to the end-Triassic mass extinction (201.6 Ma) on land, decimating all medium- to large-sized nondinosaurian, noninsulated continental reptiles. In contrast, insulated dinosaurs were already well adapted to cold temperatures, and not only survived but also underwent a rapid adaptive radiation and ecological expansion in the Jurassic, taking over regions formerly dominated by large noninsulated reptiles.

Braincase anatomy of extant Crocodylia, with new insights into the development and evolution of the neurocranium in crocodylomorphs.

Present-day crocodylians exhibit a remarkably akinetic skull with a highly modified braincase. We present a comprehensive description of the neurocranial osteology of extant crocodylians, with notes on the development of individual skeletal elements and a discussion of the terminology used for this project. The quadrate is rigidly fixed by multiple contacts with most braincase elements. The parabasisphenoid is sutured to the pterygoids (palate) and the quadrate (suspensorium); as a result, the basipterygoid joint is completely immobilized. The prootic is reduced and externally concealed by the quadrate. It has a verticalized buttress that participates in the canal for the temporal vasculature. The ventrolateral processes of the otoccipitals completely cover the posteroventral region of the braincase, enclose the occipital nerves and blood vessels in narrow bony canals and also provide additional sutural contacts between the braincase elements and further consolidate the posterior portion of the crocodylian skull. The otic capsule of crocodylians has a characteristic cochlear prominence that corresponds to the lateral route of the perilymphatic sac. Complex internal structures of the otoccipital (extracapsular buttress) additionally arrange the neurovascular structures of the periotic space of the cranium. Most of the braincase elements of crocodylians are excavated by the paratympanic pneumatic sinuses. The braincase in various extant crocodylians has an overall similar structure with some consistent variation between taxa. Several newly observed features of the braincase are present in Gavialis gangeticus and extant members of Crocodylidae to the exclusion of alligatorids: the reduced exposure of the prootic buttress on the floor of the temporal canal, the sagittal nuchal crest of the supraoccipital projecting posteriorly beyond the postoccipital processes and the reduced paratympanic pneumaticity. The most distinctive features of the crocodylian braincase (fixed quadrate and basipterygoid joint, consolidated occiput) evolved relatively rapidly at the base of Crocodylomorpha and accompanied the initial diversification of this clade during the Late Triassic and Early Jurassic. We hypothesize that profound rearrangements in the individual development of the braincases of basal crocodylomorphs underlie these rapid evolutionary modifications. These rearrangements are likely reflected in the embryonic development of extant crocodylians and include the involvement of neomorphic dermal anlagen in different portions of the developing chondrocranium, the extensive ossification of the palatoquadrate cartilage as a single expanded quadrate and the anteromedial inclination of the quadrate.

Osteology, relationships and functional morphology of Weigeltisaurus jaekeli (Diapsida, Weigeltisauridae) based on a complete skeleton from the Upper Permian Kupferschiefer of Germany.

BACKGROUND: Weigeltisauridae is a clade of small-bodied diapsids characterized by a horned cranial frill, slender trunk and limbs, and a patagium supported by elongated bony rods. Partial skeletons and fragments are definitively known only from upper Permian (Lopingian) rocks in England, Germany, Madagascar and Russia. Despite these discoveries, there have been few detailed descriptions of weigeltisaurid skeletons, and the homologies of many skeletal elements-especially the rods supporting the patagium-remain the subject of controversy. MATERIALS & METHODS: Here, we provide a detailed description of a nearly complete skeleton of Weigeltisaurus jaekeli from the upper Permian (Lopingian: Wuchiapingian) Kupferschiefer of Lower Saxony, Germany. Briefly addressed by past authors, the skeleton preserves a nearly complete skull, postcranial axial skeleton, appendicular skeleton, and patagial supports. Through comparisons with extant and fossil diapsids, we examine the hypotheses for the homologies of the patagial rods. To examine the phylogenetic position of Weigeltisauridae and characterize the morphology of the clade, we integrate the material and other weigeltisaurids into a parsimony-based phylogenetic analysis focused on Permo-Triassic non-saurian Diapsida and early Sauria (61 taxa, 339 characters). RESULTS: We recognize a number of intriguing anatomical features in the weigeltisaurid skeleton described here, including hollow horns on the post-temporal arch, lanceolate teeth in the posterior portion of the maxilla, the absence of a bony arch connecting the postorbital and squamosal bones, elongate and slender phalanges that resemble those of extant arboreal squamates, and patagial rods that are positioned superficial to the lateral one third of the gastral basket. Our phylogenetic study recovers a monophyletic Weigeltisauridae including Coelurosauravus elivensis, Weigeltisaurus jaekeli, and Rautiania spp. The clade is recovered as the sister taxon to Drepanosauromorpha outside of Sauria (=Lepidosauria + Archosauria). CONCLUSIONS: Our anatomical observations and phylogenetic analysis show variety of plesiomorphic diapsid characters and apomorphies of Weigeltisauridae in the specimen described here. We corroborate the hypothesis that the patagial ossifications are dermal bones unrelated to the axial skeleton. The gliding apparatus of weigeltisaurids was constructed from dermal elements unknown in other known gliding diapsids. SMNK-PAL 2882 and other weigeltisaurid specimens highlight the high morphological disparity of Paleozoic diapsids already prior to their radiation in the early Mesozoic.

First rebbachisaurid sauropod dinosaur from Asia.

Dzharatitanis kingi gen. et sp. nov. is based on an isolated anterior caudal vertebra (USNM 538127) from the Upper Cretaceous (Turonian) Bissekty Formation at Dzharakuduk, Uzbekistan. Phylogenetic analysis places the new taxon within the diplodocoid clade Rebbachisauridae. This is the first rebbachisaurid reported from Asia and one of the youngest rebbachisaurids in the known fossil record. The caudal is characterized by a slightly opisthocoelous centrum, 'wing-like' transverse processes with large but shallow PRCDF and POCDF, and the absence of a hyposphenal ridge and of TPRL and TPOL. The neural spine has high SPRL, SPDL, SPOL, and POSL and is pneumatized. The apex of neural spine is transversely expanded and bears triangular lateral processes. The new taxon shares with Demandasaurus and the Wessex rebbachisaurid a high SPDL on the lateral side of the neural spine, separated from SPRL and SPOL. This possibly suggests derivation of Dzharatitanis from European rebbachisaurids. This is the second sauropod group identified in the assemblage of non-avian dinosaurs from the Bissekty Formation, in addition to a previously identified indeterminate titanosaurian.

A new non-mammalian eucynodont from the Chinle Formation (Triassic: Norian), and implications for the early Mesozoic equatorial cynodont record.

The Upper Triassic tetrapod fossil record of North America features a pronounced discrepancy between the assemblages of present-day Virginia and North Carolina relative to those of the American Southwest. While both are typified by large-bodied archosaurian reptiles like phytosaurs and aetosaurs, the latter notably lacks substantial representation of mammal relatives, including cynodonts. Recently collected non-mammalian eucynodontian jaws from the middle Norian Blue Mesa Member of the Chinle Formation in northeastern Arizona shed light on the Triassic cynodont record from western equatorial Pangaea. Importantly, they reveal new biogeographic connections to eastern equatorial Pangaea as well as southern portions of the supercontinent. This discovery indicates that the faunal dissimilarity previously recognized between the western and eastern portions of equatorial Pangaea is overstated and possibly reflects longstanding sampling biases, rather than a true biogeographic pattern.

Colobops: a juvenile rhynchocephalian reptile (Lepidosauromorpha), not a diminutive archosauromorph with an unusually strong bite.

Correctly identifying taxa at the root of major clades or the oldest clade-representatives is critical for meaningful interpretations of evolution. A small, partially crushed skull from the Late Triassic (Norian) of Connecticut, USA, originally described as an indeterminate rhynchocephalian saurian, was recently named Colobops noviportensis and reinterpreted as sister to all remaining Rhynchosauria, one of the earliest and globally distributed groups of herbivorous reptiles. It was also interpreted as having an exceptionally reinforced snout and powerful bite based on an especially large supratemporal fenestra. Here, after a re-analysis of the original scan data, we show that the skull was strongly dorsoventrally compressed post-mortem, with most bones out of life position. The cranial anatomy is consistent with that of other rhynchocephalian lepidosauromorphs, not rhynchosaurs. The 'reinforced snout' region and the 'exceptionally enlarged temporal region' are preservational artefacts and not exceptional among clevosaurid rhynchocephalians. Colobops is thus not a key taxon for understanding diapsid feeding apparatus evolution.

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.

Palaeontology: Many Babies or Bigger Brains?

A cynodont from the Early Jurassic reveals unexpectedly large litters in these early mammal relatives, supporting the hypothesis that an increase in brain size was connected to smaller litters during the early evolution of mammals.

The oldest record of Alvarezsauridae (Dinosauria: Theropoda) in the Northern Hemisphere.

Procoelous caudal vertebrae, a carpometacarpus with a hypertrophied metacarpal II, and robust proximal and ungual phalanges of manual digit II of a small theropod dinosaur from the Upper Cretaceous (Turonian) Bissekty Formation at Dzharakuduk, Uzbekistan, show unequivocal synapomorphies of the clade Alvarezsauridae and thus are referred to it. The caudal vertebrae have a unique longitudinal canal within the neural arch. The carpometacarpus, with metacarpal III occupying about one third of the width of the carpometacarpus, shows the most plesiomorphic stage of the evolution of the forelimb among known alvarezsaurids. The proximal phalanx of manual digit II differs from the corresponding bone in Parvicursorinae in having a less asymmetrical proximal articular surface without a dorsal process and short ventral ridges. The ungual phalanx of manual digit II has laterally open ventral foramina. The Bissekty alvarezsaurid possibly represents a basal parvicursorine and is the stratigraphically oldest known alvarezsaurid in Asia known to date.

New tyrannosaur from the mid-Cretaceous of Uzbekistan clarifies evolution of giant body sizes and advanced senses in tyrant dinosaurs.

Tyrannosaurids--the familiar group of carnivorous dinosaurs including Tyrannosaurus and Albertosaurus--were the apex predators in continental ecosystems in Asia and North America during the latest Cretaceous (ca. 80-66 million years ago). Their colossal sizes and keen senses are considered key to their evolutionary and ecological success, but little is known about how these features developed as tyrannosaurids evolved from smaller basal tyrannosauroids that first appeared in the fossil record in the Middle Jurassic (ca. 170 million years ago). This is largely because of a frustrating 20+ million-year gap in the mid-Cretaceous fossil record, when tyrannosauroids transitioned from small-bodied hunters to gigantic apex predators but from which no diagnostic specimens are known. We describe the first distinct tyrannosauroid species from this gap, based on a highly derived braincase and a variety of other skeletal elements from the Turonian (ca. 90-92 million years ago) of Uzbekistan. This taxon is phylogenetically intermediate between the oldest basal tyrannosauroids and the latest Cretaceous forms. It had yet to develop the giant size and extensive cranial pneumaticity of T. rex and kin but does possess the highly derived brain and inner ear characteristic of the latest Cretaceous species. Tyrannosauroids apparently developed huge size rapidly during the latest Cretaceous, and their success in the top predator role may have been enabled by their brain and keen senses that first evolved at smaller body size.

The diapsid origin of turtles.

The origin of turtles has been a persistent unresolved problem involving unsettled questions in embryology, morphology, and paleontology. New fossil taxa from the early Late Triassic of China (Odontochelys) and the Late Middle Triassic of Germany (Pappochelys) now add to the understanding of (i) the evolutionary origin of the turtle shell, (ii) the ancestral structural pattern of the turtle skull, and (iii) the phylogenetic position of Testudines. As has long been postulated on the basis of molecular data, turtles evolved from diapsid reptiles and are more closely related to extant diapsids than to parareptiles, which had been suggested as stem group by some paleontologists. The turtle cranium with its secondarily closed temporal region represents a derived rather than a primitive condition and the plastron partially evolved through the fusion of gastralia.

The furculae of the dromaeosaurid dinosaur Dakotaraptor steini are trionychid turtle entoplastra.

Dakotaraptor steini is a recently described dromaeosaurid dinosaur from the Upper Cretaceous (Maastrichtian) Hell Creek Formation of South Dakota. Included within the D. steini hypodigm are three elements originally identified as furculae, one of which was made part of the holotype specimen. We show that the elements described as D. steini 'furculae' are not theropod dinosaur furculae, but are rather trionychid turtle entoplastra referable to cf. Axestemys splendida. The hypodigm of D. steini should be adjusted accordingly.

Anatomy of the Enigmatic Reptile Elachistosuchus huenei Janensch, 1949 (Reptilia: Diapsida) from the Upper Triassic of Germany and Its Relevance for the Origin of Sauria.

The holotype and only known specimen of the enigmatic small reptile Elachistosuchus huenei Janensch, 1949 from the Upper Triassic (Norian) Arnstadt Formation of Saxony-Anhalt (Germany) is redescribed using µCT scans of the material. This re-examination revealed new information on the morphology of this taxon, including previously unknown parts of the skeleton such as the palate, braincase, and shoulder girdle. Elachistosuchus is diagnosed especially by the presence of the posterolateral process of the frontal, the extension of the maxillary tooth row to the posterior margin of the orbit, the free posterior process of the jugal, and the notched anterior margin of the interclavicle. Phylogenetic analyses using two recently published character-taxon matrices recovered conflicting results for the phylogenetic position of Elachistosuchus-either as an archosauromorph, as a lepidosauromorph or as a more basal, non-saurian diapsid. These different placements highlight the need of a thorough revision of critical taxa and new character sets used for inferring neodiapsid relationships.

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.