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Psittacosaurus is one of the most abundant and speciose genera in the Dinosauria, with fifteen named species. The genus is geographically and temporally widespread with large sample sizes of several of the nominal species allowing detailed analysis of intra- and interspecific variation. We present a reanalysis of three separate, coeval species within the Psittacosauridae; P. lujiatunensis, P. major, and Hongshanosaurus houi from the Lujiatun beds of the Yixian Formation, northeastern China, using three-dimensional geometric morphometrics on a sample set of thirty skulls in combination with a reevaluation of the proposed character states for each species. Using these complementary methods, we show that individual and taphonomic variation are the joint causes of a large range of variation among the skulls when they are plotted in a morphospace. Our results demonstrate that there is only one species of Psittacosaurus within the Lujiatun beds and that the three nominal species represent different taphomorphotypes of P. lujiatunensis. The wide range of geometric morphometric variation in a single species of Psittacosaurus implies that the range of variation found in other dinosaurian groups may also be related to taphonomic distortion rather than interspecific variation. As the morphospace is driven primarily by variation resulting from taphonomic distortion, this study demonstrates that the geometric morphometric approach can only be used with great caution to delineate interspecific variation in Psittacosaurus and likely other dinosaur groups without a complementary evaluation of character states. This study presents the first application of 3D geometric morphometrics to the dinosaurian morphospace and the first attempt to quantify taphonomic variation in dinosaur skulls.

Living tetrapods owe their existence to a critical moment 360–340 million years ago when their ancestors walked on land. Vertebrae are central to locomotion, yet systematic testing of correlations between vertebral form and terrestriality and subsequent reinvasions of aquatic habitats is lacking, obscuring our understanding of movement capabilities in early tetrapods. Here, we quantified vertebral shape across a diverse group of Paleozoic amphibians (Temnospondyli) encompassing different habitats and nearly the full range of early tetrapod vertebral shapes. We demonstrate that temnospondyls were likely ancestrally terrestrial and had several early reinvasions of aquatic habitats. We find a greater diversity in temnospondyl vertebrae than previously known. We also overturn long-held hypotheses centered on weight-bearing, showing that neural arch features, including muscle attachment, were plastic across the water-land divide and do not provide a clear signal of habitat preferences. In contrast, intercentra traits were critical, with temnospondyls repeatedly converging on distinct forms in terrestrial and aquatic taxa, with little overlap between. Through our geometric morphometric study, we have been able to document associations between vertebral shape and environmental preferences in Paleozoic tetrapods and to reveal morphological constraints imposed by vertebrae to locomotion, independent of ancestry.

Despite current interest in estimating the diversity of fossil and extant groups, little effort has been devoted to estimating the diversity of dinosaurs. Here we estimate the diversity of nonavian dinosaurs at ≈1,850 genera, including those that remain to be discovered. With 527 genera currently described, at least 71% of dinosaur genera thus remain unknown. Although known diversity declined in the last stage of the Cretaceous, estimated diversity was steady, suggesting that dinosaurs as a whole were not in decline in the 10 million years before their ultimate extinction. We also show that known diversity is biased by the availability of fossiliferous rock outcrop. Finally, by using a logistic model, we predict that 75% of discoverable genera will be known within 60-100 years and 90% within 100-140 years. Because of nonrandom factors affecting the process of fossil discovery (which preclude the possibility of computing realistic confidence bounds), our estimate of diversity is likely to be a lower bound.

Eolambia caroljonesa is known from copious remains from the lower Cenomanian Mussentuchit Member of the Cedar Mountain Formation in eastern Utah; however, the taxon has been only briefly described. Thus, we present herein a complete osteological description of Eolambia. The description of Eolambia presented here is based upon the holotype partial skeleton (CEUM 9758), paratype partial skull (CEUM 5212), and abundant disarticulated elements from two bonebeds that contain juvenile individuals. These remains allow the skeletal anatomy of Eolambia to be documented almost fully and a revised diagnosis to be proposed. The description provided here facilitates comparisons between Eolambia and other iguanodontians and allows Eolambia to be coded for additional characters in phylogenetic analyses. The close affinity between Eolambia and Probactrosaurus gobiensis from the Early Cretaceous of China supports previous hypotheses of faunal interchange between Asia and North America in the early Late Cretaceous.

There is little consistency in the notation and orientation terminology used in discussions of non-mammalian fossil venebrate dentitions. The standardization of this terminology, as done in the medical and dental sciences, would facilitate all future research on fossil teeth. For mammals, we recommend following convention, where incisors, canines, premolare, and molars are abbreviated as In, Cn, Pn, and Mn (n = tooth number) in upper jaws and as in, cn, pn, and mn in lower jaws. Right, left, and deciduous teeth are indicated by R, L, and D (e.g., DP4, Rp2). For non-mammals, which can have dentigerous premaxillae, maxillae, and dentaries, as well as additional tooth-bearing bones (e.g., vomers, palatines, pterygoids, ectopterygoids, sphenoids, splenials, and even parasphenoids), we encourage identifying teeth using the bone abbreviation (e.g., pmn, mxn, dn, vn, pain). A number and slash (/) combination can be used to distinguish between multiple tooth rows (e.g., Pal1/n, Pal2/n), and specimen-specific maps can be created for very complicated dentitions. We suggest the use of the terms mesial and distal to designate tooth surfaces and directions facing toward and away from the mandibular symphysis. Labial is offered for those surfaces and directions facing the lips or cheeks and lingual for those facing the tongue. We offer the terms basal for the direction toward crown bases, apical for the direction toward crown tips, occlusal for views of the occlusal surfaces, and basal and root apical for views of crown bases and roots, respectively.

Increased excavation of dinosaurs from China over the last two decades has enriched the record of Asian titanosauriform sauropods. However, the relationships of these sauropods remain contentious, and hinges on a few well-preserved taxa, such as Euhelopus zdanskyi. Here we describe a new sauropod, Yongjinglong datangi gen. nov. et sp. nov., from the Lower Cretaceous Hekou Group in the Lanzhou Basin of Gansu Province, northwestern China. Yongjinglong datangi is characterized by the following unique combination of characters, including seven autapomorphies: long-crowned, spoon-shaped premaxillary tooth; axially elongate parapophyses on the cervical vertebra; very deep lateral pneumatic foramina on the lateral surfaces of the cervical and cranial dorsal vertebral centra; low, unbifurcated neural spine fused with the postzygapophyses to form a cranially-pointing, triangular plate in a middle dorsal vertebra; an \"XI\"-shaped configuration of the laminae on the arches of the middle dorsal vertebrae; a very long scapular blade with straight cranial and caudal edges; and a tall, deep groove on the lateral surface of the distal shaft of the radius. The new specimen shares several features with other sauropods: a pronounced M. triceps longus tubercle on the scapula and ventrolaterally elongated parapophyses in its cervical vertebra as in Euhelopodidae. Based on phylogenetic analyses Yongjinglong datangi is highly derived within Titanosauria, which suggests either a remarkable convergence with more basal titanosauriform sauropods in the Early Cretaceous or a retention of plesiomorphic features that were lost in other titanosaurians. The morphology and remarkable length of the scapulocoracoid reveal an unusual relationship between the shoulder and the middle trunk: the scapulocoracoid spans over half of the length of the trunk. The medial, notch-shaped coracoid foramen and the partially fused scapulocoracoid synostosis suggest that the specimen is a subadult individual. This specimen sheds new light on the diversity of Early Cretaceous Titanosauriformes in China.

Nothronychus was the first definitive therizinosaurian discovered in North America and currently represents the most specialized North American therizinosaurian genus. It is known from two species, No. mckinleyi from the Moreno Hill Formation (middle Turonian) in west-central New Mexico, and No. graffami from the Tropic Shale (early Turonian) in south-central Utah. Both species are represented by partial to nearly complete skeletons that have helped elucidate evolutionary trends in Therizinosauria. In spite of the biogeographical and evolutionary importance of these two taxa, neither has received a detailed description. Here, we present comprehensive descriptions of No. mckinleyi and No. graffami, the latter of which represents the most complete therizinosaurid skeleton known to date. We amend previous preliminary descriptions of No. mckinleyi and No. graffami based on these new data and modify previous character states based on an in-depth morphological analysis. Additionally, we review the depositional history of both specimens of Nothronychus and compare their taphonomic modes. We demonstrate that the species were not only separated geographically, but also temporally. Based on ammonoid biozones, the species appear to have been separated by at least 1.5 million years and up to 3 million years. We then discuss the impacts of diagenetic deformation on morphology and reevaluate potentially diagnostic characters in light of these new data. For example, the ulna of No. mckinleyi is curved whereas the ulna of No. graffami was considered straight, a character originally separating the two species. However, here we present the difference as much more likely related to diagenetic compression in No. graffami rather than as a true biologic difference. Finally, we include copies of three-dimensional surface scans of all major bones for both taxa for reference.

Terrestrial tetrapods use their claws to interact with their environments in a plethora of ways. Birds in particular have developed a diversity of claw shapes since they are often not bound to terrestrial locomotion and have heterogeneous body masses ranging several orders of magnitude. Numerous previous studies have hypothesized a connection between pedal claw shape and ecological mode in birds, yet have generated conflicting results, spanning from clear ecological groupings based on claw shape to a complete overlap of ecological modes. The majority of these studies have relied on traditional morphometric arc measurements of keratinous sheaths and have variably accounted for likely confounding factors such as body mass and phylogenetic relatedness. To better address the hypothesized relationship between ecology and claw shape in birds, we collected 580 radiographs allowing visualization of the bony core and keratinous sheath shape in 21 avian orders. Geometric morphometrics was used to quantify bony core and keratinous sheath shape and was compared to results using traditional arc measurements. Neither approach significantly separates bird claws into coarse ecological categories after integrating body size and phylogenetic relatedness; however, some separation between ecological groups is evident and we find a gradual shift from the claw shape of ground‐dwelling birds to those of predatory birds. Further, the bony claw core and keratinous sheath are significantly correlated, and the degree of functional integration does not differ across ecological groups. Therefore, it is likely possible to compare fossil bony cores with extant keratinous sheaths after applying corrections. Finally, traditional metrics and geometric morphometric shape are significantly, yet loosely correlated. Based on these results, future workers are encouraged to use geometric morphometric approaches to study claw geometry and account for confounding factors such as body size, phylogeny, and individual variation prior to predicting ecology in fossil taxa. Claw shape was quantified using geometric morphometrics and was compared to results using traditional arc measurements. Neither traditional nor geometric morphometrics are capable of significantly separating bird claws into coarse ecological categories after integrating body size and phylogenetic relatedness. Based on these results, future workers are encouraged to use geometric morphometric approaches to study claw geometry and account for confounding factors such as body size, phylogeny, and individual variation prior to predicting ecology in fossil taxa.

While centrosaurines and ceratopsids in general are abundant in the Late Campanian of northern Laramidia, they are much less commonly found in southern Laramidia. This has supported hypotheses of dinosaur provinciality and endemism in the Late Cretaceous with the delineation of at least two separate faunal zones, north and south Laramidia. There have been 12 genera of centrosaurines recognized from northern Laramidia while two genera, Diabloceratops and Nasutoceratops, have been named from southern Laramidia. We present an osteological description and taphonomic outline for a new centrosaurine ceratopsid from the Aguja Formation of northern Coahuila, Mexico that is not currently diagnosable to the generic level, but likely represents a new taxon. Further, we have included three-dimensional surface scans of all material attributed to this animal. Considering the large number of centrosaurines from northern Laramidia, it is likely that cladistic analyses are biased towards this faunal zone. New findings of southern centrosaurines are needed to correct this bias. This discovery expands the range of centrosaurines south to Coahuila, Mexico and adds new information to better characterize the morphology and taxonomy of centrosaurines from southern Laramidia and their evolution in comparison to their northern counterparts.