Possible eucynodont (Synapsida: Cynodontia) tracks from a lacustrine facies in the Lower Jurassic Moenave Formation of southwestern Utah.

Eight fossil tetrapod footprints from lake-shore deposits in the Lower Jurassic Moenave Formation at the St. George Dinosaur Discovery Site (SGDS) in southwestern Utah cannot be assigned to the prevalent dinosaurian (Anomoepus, Eubrontes, Gigandipus, Grallator, Kayentapus) or crocodyliform (Batrachopus) ichnotaxa at the site. The tridactyl and tetradactyl footprints are incomplete, consisting of digit- and digit-tip-only imprints. Seven of the eight are likely pes prints; the remaining specimen is a possible manus print. The pes prints have digit imprint morphologies and similar anterior projections and divarication angles to those of Brasilichnium, an ichnotaxon found primarily in eolian paleoenvironments attributed to eucynodont synapsids. Although their incompleteness prevents clear referral to Brasilichnium, the SGDS tracks nevertheless suggest a eucynodont track maker and thus represent a rare, Early Mesozoic occurrence of such tracks outside of an eolian paleoenvironment.

Morphology and distribution of scales, dermal ossifications, and other non-feather integumentary structures in non-avialan theropod dinosaurs.

Modern birds are typified by the presence of feathers, complex evolutionary innovations that were already widespread in the group of theropod dinosaurs (Maniraptoriformes) that include crown Aves. Squamous or scaly reptilian-like skin is, however, considered the plesiomorphic condition for theropods and dinosaurs more broadly. Here, we review the morphology and distribution of non-feathered integumentary structures in non-avialan theropods, covering squamous skin and naked skin as well as dermal ossifications. The integumentary record of non-averostran theropods is limited to tracks, which ubiquitously show a covering of tiny reticulate scales on the plantar surface of the pes. This is consistent also with younger averostran body fossils, which confirm an arthral arrangement of the digital pads. Among averostrans, squamous skin is confirmed in Ceratosauria (Carnotaurus), Allosauroidea (Allosaurus, Concavenator, Lourinhanosaurus), Compsognathidae (Juravenator), and Tyrannosauroidea (Santanaraptor, Albertosaurus, Daspletosaurus, Gorgosaurus, Tarbosaurus, Tyrannosaurus), whereas dermal ossifications consisting of sagittate and mosaic osteoderms are restricted to Ceratosaurus. Naked, non-scale bearing skin is found in the contentious tetanuran Sciurumimus, ornithomimosaurians (Ornithomimus) and possibly tyrannosauroids (Santanaraptor), and also on the patagia of scansoriopterygids (Ambopteryx, Yi). Scales are surprisingly conservative among non-avialan theropods compared to some dinosaurian groups (e.g. hadrosaurids); however, the limited preservation of tegument on most specimens hinders further interrogation. Scale patterns vary among and/or within body regions in Carnotaurus, Concavenator and Juravenator, and include polarised, snake-like ventral scales on the tail of the latter two genera. Unusual but more uniformly distributed patterning also occurs in Tyrannosaurus, whereas feature scales are present only in Albertosaurus and Carnotaurus. Few theropods currently show compelling evidence for the co-occurrence of scales and feathers (e.g. Juravenator, Sinornithosaurus), although reticulate scales were probably retained on the mani and pedes of many theropods with a heavy plumage. Feathers and filamentous structures appear to have replaced widespread scaly integuments in maniraptorans. Theropod skin, and that of dinosaurs more broadly, remains a virtually untapped area of study and the appropriation of commonly used techniques in other palaeontological fields to the study of skin holds great promise for future insights into the biology, taphonomy and relationships of these extinct animals.

A trace fossil made by a walking crayfish or crayfish-like arthropod from the Lower Jurassic Moenave Formation of southwestern Utah, USA.

New invertebrate trace fossils from the Lower Jurassic Moenave Formation at the St. George Dinosaur Discovery Site at Johnson Farm (SGDS) continue to expand the ichnofauna at the site. A previously unstudied arthropod locomotory trace, SGDS 1290, comprises two widely spaced, thick, gently undulating paramedial impressions flanked externally by small, tapered to elongate tracks with a staggered to alternating arrangement. The specimen is not a variant of any existing ichnospecies, but bears a striking resemblance to modern, experimentally generated crayfish walking traces, suggesting a crayfish or crayfish-like maker for the fossil. Because of its uniqueness, we place it in a new ichnospecies, Siskemia eurypyge. It is the first fossil crayfish or crayfish-like locomotion trace ever recorded.

Acherontiscus caledoniae: the earliest heterodont and durophagous tetrapod.

The enigmatic tetrapod Acherontiscus caledoniae from the Pendleian stage of the Early Carboniferous shows heterodontous and durophagous teeth, representing the earliest known examples of significant adaptations in tetrapod dental morphology. Tetrapods of the Late Devonian and Early Carboniferous (Mississippian), now known in some depth, are generally conservative in their dentition and body morphologies. Their teeth are simple and uniform, being cone-like and sometimes recurved at the tip. Modifications such as keels occur for the first time in Early Carboniferous Tournaisian tetrapods. Acherontiscus, dated as from the Pendleian stage, is notable for being very small with a skull length of about 15 mm, having an elongate vertebral column and being limbless. Cladistic analysis places it close to the Early Carboniferous adelospondyls, aïstopods and colosteids and supports the hypothesis of 'lepospondyl' polyphyly. Heterodonty is associated with a varied diet in tetrapods, while durophagy suggests a diet that includes hard tissue such as chitin or shells. The mid-Carboniferous saw a significant increase in morphological innovation among tetrapods, with an expanded diversity of body forms, skull shapes and dentitions appearing for the first time.

Carboniferous-Permian climate change constrained early land vertebrate radiations.

The Carboniferous-Permian transition (CPT) was Earth's last pre-Quaternary icehouse-greenhouse transition, recording major shifts in late Palaeozoic climate regimes and increased continental seasonality over approximately 40 Myr. Its parallels to Quaternary climate change have inspired recent investigations into the impacts of purported rainforest collapse on palaeotropical vertebrate diversity, but little is known about how the protracted spatial dynamics of this transition impacted the emergence of modern tetrapod lineages. Here, we apply ecological ordinance analyses on a dataset of 286 CPT fossil vertebrate localities binned across four physiographic regions forming a palaeoequatorial transect. Our results clarify the spatiotemporal expansion of land-living vertebrates, demonstrating that the reduction of tropical wetlands accommodated emerging dryland-adapted amniote faunas from a western Pangaean epicentre. We call this west-east lag the 'Vaughn-Olson model': CPT climatic transitions were regionally diachronous with delayed proliferation of amniote-dominated dryland assemblages in the east. By combining our ecological analyses with a phylogenetic approach, we demonstrate that this pattern also applies to some co-occurring total-group amphibians, suggesting that there was pervasive selection for such dryland adaptations across the crown tetrapod tree, in contrast with stem tetrapods and 'fishes'.

New basal iguanodonts from the Cedar Mountain formation of Utah and the evolution of thumb-spiked dinosaurs.

BACKGROUND: Basal iguanodontian dinosaurs were extremely successful animals, found in great abundance and diversity almost worldwide during the Early Cretaceous. In contrast to Europe and Asia, the North American record of Early Cretaceous basal iguanodonts has until recently been limited largely to skulls and skeletons of Tenontosaurus tilletti. METHODOLOGY/PRINCIPAL FINDINGS: Herein we describe two new basal iguanodonts from the Yellow Cat Member of the Cedar Mountain Formation of eastern Utah, each known from a partial skull and skeleton. Iguanacolossus fortis gen. et sp. nov. and Hippodraco scutodens gen. et sp. nov. are each diagnosed by a single autapomorphy and a unique combination of characters. CONCLUSIONS/SIGNIFICANCE: Iguanacolossus and Hippodraco add greatly to our knowledge of North American basal iguanodonts and prompt a new comprehensive phylogenetic analysis of basal iguanodont relationships. This analysis indicates that North American Early Cretaceous basal iguanodonts are more basal than their contemporaries in Europe and Asia.

Bird-like anatomy, posture, and behavior revealed by an early jurassic theropod dinosaur resting trace.

BACKGROUND: Fossil tracks made by non-avian theropod dinosaurs commonly reflect the habitual bipedal stance retained in living birds. Only rarely-captured behaviors, such as crouching, might create impressions made by the hands. Such tracks provide valuable information concerning the often poorly understood functional morphology of the early theropod forelimb. METHODOLOGY/PRINCIPAL FINDINGS: Here we describe a well-preserved theropod trackway in a Lower Jurassic ( approximately 198 million-year-old) lacustrine beach sandstone in the Whitmore Point Member of the Moenave Formation in southwestern Utah. The trackway consists of prints of typical morphology, intermittent tail drags and, unusually, traces made by the animal resting on the substrate in a posture very similar to modern birds. The resting trace includes symmetrical pes impressions and well-defined impressions made by both hands, the tail, and the ischial callosity. CONCLUSIONS/SIGNIFICANCE: The manus impressions corroborate that early theropods, like later birds, held their palms facing medially, in contrast to manus prints previously attributed to theropods that have forward-pointing digits. Both the symmetrical resting posture and the medially-facing palms therefore evolved by the Early Jurassic, much earlier in the theropod lineage than previously recognized, and may characterize all theropods.