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Abstract Silesaurids (Archosauria: Dinosauriformes) are found in Middle to Upper Triassic deposits across Pangea, but few stratigraphic sections record the evolution of the group in one geographic area over millions of years. Here, we describe silesaurid remains from the oldest of the Upper Triassic stratigraphic sequence from the base of the Dockum Group, from the type locality of the Otischalkian faunachronozone. Isolated limb bones diagnostic of silesaurids include humeri, femora, and tibiae of a seemingly uniqueSilesaurus‐like taxon from the same locality (Otis Chalk Quarry 3). The femora consist of four specimens of different lengths that sample the variation of character states associated with ontogeny, also sampled previously in both silesaurids (e.g.,Asilisaurus kongweandSilesaurus opolensis) and within neotheropods within Dinosauria (e.g.,Coelophysis bauri). Our observations of the variation in the silesaurid sample further reinforce the interpretation of high variation of morphological features common in dinosauriforms. Furthermore, we show that overpreparation of bone surfaces has hidden some of this variation in previous interpretations. The tibia growth series shows that the fibular crest of the tibia develops during ontogeny, yet another phylogenetically informative character for dinosaurs and their kin that is at least ontogenetically variable in silesaurids. The presence of silesaurids at the base of the Dockum Group (late Carnian or early Norian) conclusively shows that the group was present near the onset of deposition of Upper Triassic rocks and survived for millions of years in the same geographic area at low latitudes throughout the Late Triassic.more » « less
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Abstract Fossils of embryonic and hatchling individuals can provide invaluable insight into the evolution of prenatal morphologies, heterochronies, and allometric trajectories within Archosauria but are exceptionally rare in the Triassic fossil record, obscuring a critical aspect of archosaurian biology during their evolutionary origins. Microvertebrate sampling at a single bonebed in the Upper Triassic Chinle Formation within Petrified Forest National Park has yielded diminutive archosauriform femora (PEFO 45274, PEFO 45199) with estimated and measured femoral lengths of ~31 mm and ~ 37 mm, respectively. These new specimens provide the unique opportunity to assess the preservation, body size, and growth dynamics of skeletally immature archosauriforms in North America and compare the growth dynamics of archosauromorphs within an evolutionary and ontogenetic context. We assign PEFO 45199 and PEFO 45274 to Phytosauria (Archosauriformes) based on their strongly sigmoidal shape in lateral view, the presence of proximal anterolateral and posteromedial tubera, the absence of an anteromedial tuber of the proximal end, a teardrop‐shaped proximal outline, and a fourth trochanter that is not confluent with the proximal head. Osteohistological analyses of PEFO 45274 reveal a cortex comprising low vascularity, parallel‐fibered bone composed of primary osteons that lacks a hatching line and any lines of arrested growth. We interpret PEFO 45274 as a slow‐growing, post‐hatching individual of less than 1 year of age. Surprisingly, osteohistology of some larger phytosaur femora implies faster growth rates in comparison to PEFO 45274 based on the occasional presence of woven bone and overall higher degrees of vascular density, suggesting the ontogenetic shift from rapid‐to‐slow growth rates might not occur simply or uniformly as expected in Phytosauria and that non‐archosaurian archosauriforms may exhibit size‐dependent histological characteristics. This study highlights the importance of including osteohistology from multiple body sizes to investigate non‐archosaurian archosauriform ancestral growth rates given the phylogenetic position of phytosaurs near the divergence of Archosauria.more » « less
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Abstract A vast array of pseudosuchian body plans evolved during the diversification of the group in the Triassic Period, but few can compare to the toothless, long‐necked, and bipedal shuvosaurids. Members of this clade possess theropod‐like character states mapped on top of more plesiomorphic pseudosuchian character states, complicating our understanding of the evolutionary history of the skeleton. One taxon in this clade,Shuvosaurus inexpectatushas been assigned to various theropod dinosaur groups based on a partial skull and referred material and its postcranium was assigned to a different taxon in Pseudosuchia. After the discovery of a skeleton of a shuvosaurid with aShuvosaurus‐like skull and a pseudosuchian postcranial skeleton, it became clearShuvosaurus inexpectatuswas a pseudosuchian. Nevertheless, a number of questions have arisen about what skeletal elements belonged toShuvosaurus inexpectatus, the identification of skull bones, and the resulting implication for pseudosuchian evolution. Here, we detail the anatomy of the skeletonShuvosaurus inexpectatusthrough a critical lens, parse out the bones that belong to the taxon or those that clearly do not or may not belong to the taxon, rediagnose the taxon based on these revisions, and compare the taxon to other archosaurs. We find thatShuvosaurus inexpectatuspossesses similar anatomy to other shuvosaurids but parts of the skeleton of the taxon clarifies the anatomy of the group given that they are preserved inShuvosaurus inexpectatusbut not in others.Shuvosaurus inexpectatusis represented by at least 14 individuals from the West Texas Post Quarry (Adamanian holochronozone) and allShuvosaurus inexpectatusskeletal material from the locality pertains to skeletally immature individuals. All of the skeletons are missing most of the neural arches, ribs, and most of the forelimb. We only recognizeShuvosaurus inexpectatusfrom the Post Quarry and all other material assigned to the taxon previously is better assigned to the broader group Shuvosauridae.more » « less
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Abstract 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.more » « less
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Abstract Archosauromorph reptiles underwent rapid lineage diversification, increases in morphological and body size disparity, and expansion into new adaptive landscapes. Several of the primary early archosauromorph clades (e.g. rhynchosaurs) are easy to differentiate from others because of their characteristic body types, whereas the more lizard‐like and carnivorous forms with long necks (e.g. tanystropheids) were historically all relegated to the groups Protorosauria or Prolacertiformes. However, it is now clear that these groups are polyphyletic and that a lizard‐like, carnivorous form is plesiomorphic for Archosauromorpha, and multiple subclades started with that body plan. Among these early forms isMalerisaurusfrom the Upper Triassic of India (M. robinsonae) and the Upper Triassic of south‐western USA (M. langstoni). In this paper, we critically re‐evaluate the genus. We find both species ofMalerisaurusas valid, and identifyMalerisaurusas an early diverging, but late‐surviving, carnivorous member of Azendohsauridae within Allokotosauria. Our histological analysis and assessment of ontogenetic changes of limb bones of small and large individuals demonstrate that the skeletons of the small forms grew slowly and became more robust through ontogeny, and that the larger recovered bones are at or near the maximum size of the taxon.MalerisaurusandMalerisaurus‐like taxa were common members of the Otischalkian–Adamanian (late Carnian to mid‐Norian) faunal assemblages from Upper Triassic strata of the south western USA, but they are absent from the younger Revueltian holochronozone. Specimens from western North America show that Allokotosauria had a near‐Pangaean distribution for much of the Middle Triassic to Late Triassic.more » « less
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Reptile feeding strategies encompass a wide variety of diets and accompanying diversity in methods for subduing prey. One such strategy, the use of venom for prey capture, is found in living reptile clades like helodermatid (beaded) lizards and some groups of snakes, and venom secreting glands are also present in some monitor lizards and iguanians. The fossil record of some of these groups shows strong evidence for venom use, and this feeding strategy also has been hypothesized for a variety of extinct reptiles (e.g., archosauromorphs, anguimorphs, and a sphenodontian). However, evidence of systems for venom delivery in extinct groups and its evolutionary origins has been scarce, especially when based on more than isolated teeth. Here, we describe a potentially venomous new reptile,Microzemiotes sonselaensisgen. et sp. nov., from a partial left dentary recovered from the Sonsela Member of the Chinle Formation (middle Norian, Upper Triassic) of northeastern Arizona, U.S.A. The three dentary teeth have apices that are distally reclined relative to their bases and the tip of the posteriormost tooth curves mesially. The teeth show subthecodont implantation and are interspaced by empty sockets that terminate above the Meckelian canal, which is dorsoventrally expanded posteriorly. Replacement tooth sockets are positioned distolingually to the active teeth as in varanid-like replacement. We identify this new specimen as a diapsid reptile based on its monocuspid teeth that lack carinae and serrations. A more exclusive phylogenetic position within Diapsida is not well supported and remains uncertain. Several features of this new taxon, such as the presence of an intramandibular septum, are shared with some anguimorph squamates; however, these likely evolved independently. The teeth of the new taxon are distinctively marked by external grooves that occur on the entire length of the crown on the labial and lingual sides, as seen in the teeth of living beaded lizards. If these grooves are functionally similar to those of beaded lizards, which use the grooves to deliver venom, this new taxon represents the oldest known reptile where venom-conducting teeth are preserved within a jaw. The teeth of the new species are anatomically distinct from and ~10x smaller than those of the only other known Late Triassic hypothesized venomous reptile,Uatchitodon, supporting venom use across multiple groups of different body size classes. This new species represents the third Late Triassic reptile species to possibly have used envenomation as a feeding (and/or defensive) strategy, adding to the small number of venomous reptiles known from the Mesozoic Era.more » « less
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Abstract Non-archosaur archosauromorphs are a paraphyletic group of diapsid reptiles that were important members of global Middle and Late Triassic continental ecosystems. Included in this group are the azendohsaurids, a clade of allokotosaurians (kuehneosaurids and Azendohsauridae + Trilophosauridae) that retain the plesiomorphic archosauromorph postcranial body plan but evolved disparate cranial features that converge on later dinosaurian anatomy, including sauropodomorph-like marginal dentition and ceratopsian-like postorbital horns. Here we describe a new malerisaurine azendohsaurid from two monodominant bonebeds in the Blue Mesa Member, Chinle Formation (Late Triassic, ca. 218–220 Ma); the first occurs at Petrified Forest National Park and preserves a minimum of eight individuals of varying sizes, and the second occurs near St. Johns, Arizona. Puercosuchus traverorum n. gen. n. sp. is a carnivorous malerisaurine that is closely related to Malerisaurus robinsonae from the Maleri Formation of India and to Malerisaurus langstoni from the Dockum Group of western Texas. Dentigerous elements from Puercosuchus traverorum n. gen. n. sp. confirm that some Late Triassic tooth morphotypes thought to represent early dinosaurs cannot be differentiated from, and likely pertain to, Puercosuchus -like malerisaurine taxa. These bonebeds from northern Arizona support the hypothesis that non-archosauriform archosauromorphs were locally diverse near the middle Norian and experienced an extinction event prior to the end-Triassic mass extinction coincidental with the Adamanian-Revueltian boundary recognized at Petrified Forest National Park. The relatively late age of this early-diverging taxon (Norian) suggests that the diversity of azendohsaurids is underrepresented in Middle and Late Triassic fossil records around the world. UUID: http://zoobank.org/e6eeefd2-a0ae-47fc-8604-9f45af8c1147 .more » « less
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