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Using morphometrics to study nonavian dinosaur fossils is a practice that predates the origin of the word "dinosaur." By the 1970s, linear morphometrics had become established as a valuable tool for analyzing intra- and interspecific variation in nonavian dinosaurs. With the advent of more recent techniques such as geometric morphometrics and more advanced statistical approaches, morphometric analyses of nonavian dinosaurs have proliferated, granting unprecedented insight into many aspects of their biology and evolution. I outline the past, present, and future of morphometrics as applied to the study of nonavian dinosaurs zeroing in on five aspects of nonavian dinosaur paleobiology where morphometrics has been widely utilized to advance our knowledge: systematics, sexual dimorphism, locomotion, macroevolution, and trackways. Morphometric methods are especially susceptible to taphonomic distortion. As such, the impact of taphonomic distortion on original fossil shape is discussed as are current and future methods for quantifying and accounting for distortion with the goal of reducing the taphonomic noise to biological signal ratio. Finally, the future of morphometrics in nonavian dinosaur paleobiology is discussed as paleobiologists move into a "virtual paleobiology" framework, whereby digital renditions of fossils are captured via methods such as photogrammetry and computed tomography. These primary data form the basis for three-dimensional (3D) geometric morphometric analyses along with a slew of other forms of analyses. These 3D specimen data form part of the extended specimen and help to democratize paleobiology, unlocking the specimen from the physical museum and making the specimen available to researchers across the world., (© 2023 American Association for Anatomy.)

The secondary evolution of quadrupedality from bipedal ancestry is a rare evolutionary transition in tetrapods yet occurred convergently at least three times within ornithischian dinosaurs. Despite convergently evolving quadrupedal gait, ornithischians exhibited variable anatomy, particularly in the forelimbs, which underwent a major functional change from assisting in foraging and feeding in bipeds to becoming principal weight-bearing components of the locomotor system in quadrupeds. Here, we use three-dimensional multi-body dynamics models to demonstrate quantitatively that different quadrupedal ornithischian clades evolved distinct forelimb musculature, particularly around the shoulder. We find that major differences in glenohumeral abduction-adduction and long axis rotation muscle leverages were key drivers of mechanical disparity, thereby refuting previous hypotheses about functional convergence in major clades. Elbow muscle leverages were also disparate across the major ornithischian lineages, although high elbow extension muscle leverages were convergent between most quadrupeds. Unlike in ornithischian hind limbs, where differences are more closely tied to functional similarity than phylogenetic relatedness, mechanical disparity in ornithischian forelimbs appears to have been shaped primarily by phylogenetic constraints. Differences in ancestral bipedal taxa within each clade may have resulted in disparate ecomorphological constraints on the evolutionary pathways driving divergence in their quadrupedal descendants.

The diversity of research performed by vertebrate paleobiologists is immense, including primary descriptions of newly discovered taxa, biomechanical analyses of extant animals, large-scale macroevolutionary analyses with hundreds of taxa, and many others. This special issue of The Anatomical Record highlights the scope of the research done by paleobiologists, with an emphasis on dinosaurs and their relatives, and is structured into three broad categories of research: (1) anatomical descriptions, (2) the biology of the dinosaurs and their relatives, and (3) macroevolutionary trends. Although these topics all fall within the purview of contemporary paleobiology, many of these topics have been studied in one form or another since the beginning of dinosaur paleobiology as a field. Rather than the topics themselves, what characterizes the modern renaissance of paleobiology is the employment of newly developed quantitative techniques to analyze the relationships between taxa and their evolutionary history. While primary anatomical descriptions remain the central pillar of dinosaur paleobiology, researchers now have a suitable baseline understanding of dinosaurian anatomy and their major evolutionary relationships. Using this baseline, they are able to ask more complex questions and refine their understanding of dinosaurian evolution. The aim of this issue is to exemplify the range of topics examined within dinosaur paleobiology and to look forward to the future of paleobiology. Anat Rec, 303:645-648, 2020. © 2020 American Association for Anatomy., (© 2020 American Association for Anatomy.)

Bone histology grants substantial insight into the growth and biology of fossil vertebrates. Many of the major non-avian dinosaurian clades have been extensively sampled for bone histologic data allowing reconstruction of their growth as well as the assessment of the evolution of growth changes along phylogenies. However, horned ceratopsians are poorly represented in paleohistologic studies. Further, the ceratopsian taxa that have been examined are unevenly sampled phylogenetically with very basal forms and highly derived forms making up the majority of studied taxa. In order to rectify this, we have histologically sampled Avaceratops from Montana and Yehuecauhceratops from northern Mexico to assess how mid-sized basal centrosaurines grew relative to more basal and derived forms. Based on results from these taxa, basal centrosaurines present a mosaic of growth characters intermediate between those seen in basal ceratopsians and more derived centrosaurines. Further, Yehuecauhceratops has many lines of arrested growth preserved, suggesting that the large number of lines of arrested growth found in a high-latitude Pachyrhinosaurus specimen may be a result of phylogeny rather than geography. Since lines of arrested growth are not preserved in long bones of many ceratopsians, especially chasmosaurines, we also histologically sampled ribs of Avaceratops and Pachyrhinosaurus. However, the largest ribs were highly remodeled obscuring lines of arrested growth, making it unlikely that rib histology will clarify growth trends in ceratopsians. These centrosaurines add to the growing ceratopsian histological database and demonstrate that basal centrosaurines grew in a manner intermediate between non-ceratopsid taxa and derived centrosaurines. Anat Rec, 303:935-948, 2020. © 2019 Wiley Periodicals, Inc., (© 2019 Wiley Periodicals, Inc.)

We describe a Psittacosaurus specimen from the Lujiatun beds of the Yixian Formation in Liaoning, China with an abnormality on its left fibula. Although a large number of Psittacosaurus specimens are known, only a single example of a pathologic Psittacosaurus has been previously noted. The specific pathology in the current specimen is believed to be a healed fibular fracture as assessed through a combination of gross morphology, microcomputed tomography (microCT), and histology data. The fracture can be identified using microCT, but the degree of remodeling and the stage of fracture repair are best determined histologically. The fracture callus is made up of radially oriented spokes of woven bone in a cartilage matrix and the original cortical bone prior to the fracture has been largely eroded. A transverse histologic section taken at the level of the fracture shows the displacement of the proximal and distal parts of the fibula. The Psittacosaurus appears to have survived the break considering the deposition of circumferential non-pathologic bone at the periosteal surface outside of the callus. The combination of gross morphological description, microCT data, and histologic data allowed for a full diagnosis of the abnormality. While some previous authors have preferred gross morphological description above other methods for assessing paleopathologies, it is evident based on this specimen that an amalgam of techniques provides greater clarity to paleopathology diagnoses. Although this Psittacosaurus lived in an environment with many predators, it was able to survive with a fracture on its hindlimb, which undoubtedly would have impacted its locomotion. Anat Rec, 299:897-906, 2016. © 2016 Wiley Periodicals, Inc., (© 2016 Wiley Periodicals, Inc.)

Psittacosaurus is one of the most abundant dinosaurs known, which allows for extensive study of its growth and form. Previous studies have evaluated growth trajectories of Psittacosaurus using bone histology. However, we present the first study of Psittacosaurus comparative juvenile histology and describe the histology of Psittacosaurus within its first year of life based on multiple sections taken from an exquisite monospecific assemblage of juveniles from the Yixian Formation in Liaoning, China. Specimens studied had femur lengths ranging from 30 to 36 mm. The five juveniles examined all have similar histologic patterns in the midshaft and epiphyseal regions showing that there is limited plasticity in bone development in juvenile Psittacosaurus and that all of the specimens in the assemblage were likely the same age. The microstructure patterns are compatible with the hypothesis that Psittacosaurus was precocial and that these juveniles were neonates. Based on comparisons with other juvenile ornithischians, juvenile Psittacosaurus had a growth rate similar to Orodromeus, slower than that of Maiasaura, Dysalotosaurus, or hadrosaurs consistent with small body size. Our results support previous studies that demonstrated that the orientation of vascular canals is likely not solely reflective of growth rate, but is also affected by underlying biomechanical, structural processes. The number of studies done on theropod and sauropodomorph histology dwarfs those of ornithischians. More studies of ornithischian histology are necessary in order to better establish phylogenetic trends in microstructure and to learn more about growth in this important clade., (© 2016 Wiley Periodicals, Inc.)

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.

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.

Theropoda was one of the most successful dinosaurian clades during the Mesozoic and has remained a dominant component of faunas throughout the Cenozoic, with nearly 10,000 extant representatives. The discovery of Archaeopteryx provides evidence that avian theropods evolved at least 155 million years ago and that more than half of the tenure of avian theropods on Earth was during the Mesozoic. Considering the major changes in niche occupation for theropods resulting from the evolution of arboreal and flight capabilities, we have analyzed forelimb and hindlimb proportions among nonmaniraptoriform theropods, nonavian maniraptoriforms, and basal avialans using reduced major axis regressions, principal components analysis, canonical variates analysis, and discriminant function analysis. Our study is the first analysis on theropod limb proportions to apply phylogenetic independent contrasts and size corrections to the data to ensure that all the data are statistically independent and amenable to statistical analyses. The three ordination analyses we performed did not show any significant groupings or deviations between nonavian theropods and Mesozoic avian forms when including all limb elements. However, the bivariate regression analyses did show some significant trends between individual elements that suggested evolutionary trends of increased forelimb length relative to hindlimb length from nonmaniraptoriform theropods to nonavian maniraptoriforms to basal avialans. The increase in disparity and divergence away from the nonavian theropod body plan is well documented within Cenozoic forms. The lack of significant groupings among Mesozoic forms when examining the entire theropod body plan concurrently suggests that nonavian theropods and avian theropods did not substantially diverge in limb proportions until the Cenozoic., (© 2014 Wiley Periodicals, Inc.)

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.