The femora of Drepanosauromorpha (Reptilia: Diapsida): Implications for the functional evolution of the thigh of Sauropsida.

The femora of diapsids have undergone morphological changes related to shifts in postural and locomotor modes, such as the transition from plesiomorphic amniote and diapsid taxa to the apomorphic conditions related to a more erect posture within Archosauriformes. One remarkable clade of Triassic diapsids is the chameleon-like Drepanosauromorpha. This group is known from numerous articulated but heavily compressed skeletons that have the potential to further inform early reptile femoral evolution. For the first time, we describe the three-dimensional osteology of the femora of Drepanosauromorpha, based on undistorted fossils from the Upper Triassic Chinle Formation and Dockum Group of North America. We identify apomorphies and a combination of character states that link these femora to those in crushed specimens of drepanosauromorphs and compare our sample with a range of amniote taxa. Several characteristics of drepanosauromorph femora, including a hemispherical proximal articular surface, prominent asymmetry in the proximodistal length of the tibial condyles, and a deep intercondylar sulcus, are plesiomorphies shared with early diapsids. The femora contrast with those of most diapsids in lacking a crest-like, distally tapering internal trochanter. They bear a ventrolaterally positioned tuberosity on the femoral shaft, resembling the fourth trochanter in Archosauriformes. The reduction of an internal trochanter parallels independent reductions in therapsids and archosauriforms. The presence of a ventrolaterally positioned trochanter is also similar to that of chameleonid squamates. Collectively, these features demonstrate a unique femoral morphology for drepanosauromorphs, and suggest an increased capacity for femoral adduction and protraction relative to most other Permo-Triassic diapsids.

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.

A tiny Triassic saurian from Connecticut and the early evolution of the diapsid feeding apparatus.

Following the Permo-Triassic Extinction, large-bodied diapsid reptiles-with a body length >1 m-rapidly expanded their ecological roles. This diversification is reflected in enormous disparity in the development of the rostrum and adductor chamber. However, it is unclear how marked the diversity of the feeding apparatus was in contemporary small-bodied diapsids. Here we describe the remarkably small skull (2.5 cm long) of a saurian reptile, Colobops noviportensis, gen. et sp. nov., from the Triassic New Haven Arkose of Connecticut, USA. The taxon possesses an exceptionally reinforced snout and strikingly expanded supratemporal fossae for adductor musculature relative to any known Mesozoic or Recent diapsid of similar size. Our phylogenetic analyses support C. noviportensis as an early diverging pan-archosaur. Colobops noviportensis reveals extraordinary disparity of the feeding apparatus in small-bodied early Mesozoic diapsids, and a suite of morphologies, functionally related to a powerful bite, unknown in any small-bodied diapsid.

A bird-like skull in a Triassic diapsid reptile increases heterogeneity of the morphological and phylogenetic radiation of Diapsida.

The Triassic Period saw the first appearance of numerous amniote lineages (e.g. Lepidosauria, Archosauria, Mammalia) that defined Mesozoic ecosystems following the end Permian Mass Extinction, as well as the first major morphological diversification of crown-group reptiles. Unfortunately, much of our understanding of this event comes from the record of large-bodied reptiles (total body length > 1 m). Here we present a new species of drepanosaurid (small-bodied, chameleon-like diapsids) from the Upper Triassic Chinle Formation of New Mexico. Using reconstructions of micro-computed tomography data, we reveal the three-dimensional skull osteology of this clade for the first time. The skull presents many archaic anatomical traits unknown in Triassic crown-group reptiles (e.g. absence of bony support for the external ear), whereas other traits (e.g. toothless rostrum, anteriorly directed orbits, inflated endocranium) resemble derived avian theropods. A phylogenetic analysis of Permo-Triassic diapsids supports the hypothesis that drepanosaurs are an archaic lineage that originated in the Permian, far removed from crown-group Reptilia. The phylogenetic position of drepanosaurids indicates the presence of archaic Permian clades among Triassic small reptile assemblages and that morphological convergence produced a remarkably bird-like skull nearly 100 Myr before one is known to have emerged in Theropoda.

The skull roof tracks the brain during the evolution and development of reptiles including birds.

Major transformations in brain size and proportions, such as the enlargement of the brain during the evolution of birds, are accompanied by profound modifications to the skull roof. However, the hypothesis of concerted evolution of shape between brain and skull roof over major phylogenetic transitions, and in particular of an ontogenetic relationship between specific regions of the brain and the skull roof, has never been formally tested. We performed 3D morphometric analyses to examine the deep history of brain and skull-roof morphology in Reptilia, focusing on changes during the well-documented transition from early reptiles through archosauromorphs, including nonavian dinosaurs, to birds. Non-avialan taxa cluster tightly together in morphospace, whereas Archaeopteryx and crown birds occupy a separate region. There is a one-to-one correspondence between the forebrain and frontal bone and the midbrain and parietal bone. Furthermore, the position of the forebrain-midbrain boundary correlates significantly with the position of the frontoparietal suture across the phylogenetic breadth of Reptilia and during the ontogeny of individual taxa. Conservation of position and identity in the skull roof is apparent, and there is no support for previous hypotheses that the avian parietal is a transformed postparietal. The correlation and apparent developmental link between regions of the brain and bony skull elements are likely to be ancestral to Tetrapoda and may be fundamental to all of Osteichthyes, coeval with the origin of the dermatocranium.

Empirical and Bayesian approaches to fossil-only divergence times: A study across three reptile clades.

Estimating divergence times on phylogenies is critical in paleontological and neontological studies. Chronostratigraphically-constrained fossils are the only direct evidence of absolute timing of species divergence. Strict temporal calibration of fossil-only phylogenies provides minimum divergence estimates, and various methods have been proposed to estimate divergences beyond these minimum values. We explore the utility of simultaneous estimation of tree topology and divergence times using BEAST tip-dating on datasets consisting only of fossils by using relaxed morphological clocks and birth-death tree priors that include serial sampling (BDSS) at a constant rate through time. We compare BEAST results to those from the traditional maximum parsimony (MP) and undated Bayesian inference (BI) methods. Three overlapping datasets were used that span 250 million years of archosauromorph evolution leading to crocodylians. The first dataset focuses on early Sauria (31 taxa, 240 chars.), the second on early Archosauria (76 taxa, 400 chars.) and the third on Crocodyliformes (101 taxa, 340 chars.). For each dataset three time-calibrated trees (timetrees) were calculated: a minimum-age timetree with node ages based on earliest occurrences in the fossil record; a 'smoothed' timetree using a range of time added to the root that is then averaged over zero-length internodes; and a tip-dated timetree. Comparisons within datasets show that the smoothed and tip-dated timetrees provide similar estimates. Only near the root node do BEAST estimates fall outside the smoothed timetree range. The BEAST model is not able to overcome limited sampling to correctly estimate divergences considerably older than sampled fossil occurrence dates. Conversely, the smoothed timetrees consistently provide node-ages far older than the strict dates or BEAST estimates for morphologically conservative sister-taxa when they sit on long ghost lineages. In this latter case, the relaxed-clock model appears to be correctly moderating the node-age estimate based on the limited morphological divergence. Topologies are generally similar across analyses, but BEAST trees for crocodyliforms differ when clades are deeply nested but contain very old taxa. It appears that the constant-rate sampling assumption of the BDSS tree prior influences topology inference by disfavoring long, unsampled branches.

Extreme Modification of the Tetrapod Forelimb in a Triassic Diapsid Reptile.

The tetrapod forelimb is one of the most versatile structures in vertebrate evolution, having been co-opted for an enormous array of functions. However, the structural relationships between the bones of the forelimb have remained largely unchanged throughout the 375 million year history of Tetrapoda, with a radius and ulna made up of elongate, paralleling shafts contacting a series of shorter carpal bones. These features are consistent across nearly all known tetrapods, suggesting that the morphospace encompassed by these taxa is limited by some sort of constraint(s). Here, we report on a series of three-dimensionally preserved fossils of the small-bodied (<1 m) Late Triassic diapsid reptile Drepanosaurus, from the Chinle Formation of New Mexico, USA, which dramatically diverge from this pattern. Along with the crushed type specimen from Italy, these specimens have a flattened, crescent-shaped ulna with a long axis perpendicular to that of the radius and hyperelongate, shaft-like carpal bones contacting the ulna that are proximodistally longer than the radius. The second digit supports a massive, hooked claw. This condition has similarities to living "hook-and-pull" digging mammals and demonstrates that specialized, modern ecological roles had developed during the Triassic Period, over 200 million years ago. The forelimb bones in Drepanosaurus represent previously unknown morphologies for a tetrapod and, thus, a dramatic expansion of known tetrapod forelimb morphospace.

The monophyly of Susisuchidae (Crocodyliformes) and its phylogenetic placement in Neosuchia.

Eusuchian crocodyliforms, which include all living crocodylians, have historically been characterized by two anatomical specializations: a ball-in-socket vertebral joint and an extensive secondary hard palate with a pterygoid-bound internal choana. The Early Cretaceous neosuchian clade Susisuchidae is typically regarded as phylogenetically near Eusuchia. The putative susisuchid Isisfordia duncani was initially described as a transitional form exhibiting incipient versions of these eusuchian traits. Here we examine aspects of the morphology of Isisfordia and comment on the morphology of its putative sister taxon Susisuchus. Our reexamination supports the notion of Isisfordia possessing transitional vertebral morphology but we present a new interpretation of its palate construction that shows it to be more plesiomorphic than previously thought. The secondary choana of Isisfordia is not pterygoid bound. Instead, long palatines expand distally lapping under the pterygoid to form the anterior border of the choana as is common among many advanced neosuchians. Incorporation of these observations into an expanded phylogenetic dataset of neosuchian crocodyliforms results in a new phylogenetic hypothesis for Susisuchidae. Isisfordia and Susisuchus form a monophyletic Susisuchidae that sits near the base of Neosuchia, and is not the sister taxon of Eusuchia.