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Compared with all other living reptiles, birds grow extremely fast and possess unusually low levels of intraspecific variation during postnatal development. It is now clear that birds inherited their high rates of growth from their dinosaurian ancestors, but the origin of the avian condition of low variation during development is poorly constrained. The most well-understood growth trajectories of later Mesozoic theropods (e.g., Tyrannosaurus, Allosaurus) show similarly low variation to birds, contrasting with higher variation in extant crocodylians. Here, we show that deep within Dinosauria, among the earliest-diverging dinosaurs, anomalously high intraspecific variation is widespread but then is lost in more derived theropods. This style of development is ancestral for dinosaurs and their closest relatives, and, surprisingly, this level of variation is far higher than in living crocodylians. Among early dinosaurs, this variation is widespread across Pangaea in the Triassic and Early Jurassic, and among early-diverging theropods (ceratosaurs), this variation is maintained for 165 million years to the end of the Cretaceous. Because the Late Triassic environment across Pangaea was volatile and heterogeneous, this variation may have contributed to the rise of dinosaurian dominance through the end of the Triassic Period.I n comparison with other reptiles, avian biology is highly unusual, characterized by "hollow" bones and postcranial skeletal pneumaticity, feathers, a unique forelimb digit formula, endothermy, and rapid growth rates. However, these peculiarities initially arose in nonavian dinosaurs (1-7) in a gradual process occurring over tens of millions of years (8,9). In addition to their extremely rapid rates of growth, avian ontogeny possesses a characteristically low level of morphological variation within a species relative to that of other reptiles. The majority of individuals in a given avian species undergoes the same morphological changes during ontogeny in the same order at similar body sizes (10, 11), whereas their closest living relatives, crocodylians, possess higher variation (10,(12)(13)(14). How and when this feature of avian biology evolved is poorly constrained.This avian style of development must have evolved after its most recent common ancestor with crocodylians but before the origin of Aves. Most studies of the ontogeny of more-derived theropods (15-17) suggest that the low levels of variation that characterize avian ontogeny were present in close nonavian relatives as well. Closer to the origin of Dinosauria, morphological variation within species is widespread (18-26), but whether this variation is the result of taxonomic diversity, ontogeny, sexual dimorphism, or simple individual variation is not clear (22,23,27). Furthermore, close dinosaurian relatives or "dinosaur precursors" (e.g., Silesaurus, Asilisaurus) possess high intraspecific variation in growth sequences [i.e., sequence polymorphism (28)], suggesting that this condition could be ancestral for Dinosauria (21).In contrast to ...

Understanding ontogenetic patterns is important in vertebrate paleontology because the assessed skeletal maturity of an individual often has implications for paleobiogeography, species synonymy, paleobiology, and body size evolution of major clades. Further, for many groups the only means of confidently determining ontogenetic status of an organism is through the destructive process of histological sampling. Although the ontogenetic patterns of Late Jurassic and Cretaceous dinosaurs are better understood, knowledge of the ontogeny of the earliest dinosaurs is relatively poor because most species-level growth series known from these groups are small (usually, maximum of n ~ 5) and incomplete. To investigate the morphological changes that occur during ontogeny in early dinosaurs, I used ontogenetic sequence analysis (OSA) to reconstruct developmental sequences of morphological changes in the postcranial ontogeny of the early theropods Coelophysis bauri and Megapnosaurus rhodesiensis, both of which are known from large sample sizes (n = 174 and 182, respectively). I found a large amount of sequence polymorphism (i.e. intraspecific variation in developmental patterns) in both taxa, and especially in C. bauri, which possesses this variation in every element analyzed. Megapnosaurus rhodesiensis is similar, but it possesses no variation in the sequence of development of ontogenetic characters in the tibia and tarsus. Despite the large amount of variation in development, many characters occur consistently earlier or later in ontogeny and could therefore be important morphological features for assessing the relative maturity of other early theropods. Additionally, there is a phylogenetic signal to the order in which homologous characters appear in ontogeny, with homologous characters appearing earlier or later in developmental sequences of early theropods and the close relatives of dinosaurs, silesaurids. Many of these morphological features are important characters for the reconstruction of archosaurian phylogeny (e.g. trochanteric shelf). Because these features vary in presence or appearance with ontogeny, these characters should be used with caution when undertaking phylogenetic analyses in these groups, since a specimen may possess certain character states owing to ontogenetic stage, not evolutionary relationships.

Morphology forms the most fundamental level of data in vertebrate palaeontology because it is through interpretations of morphology that taxa are identified, creating the basis for broad evolutionary and palaeobiological hypotheses. Assessing maturity is one of the most basic aspects of morphological interpretation and provides the means to study the evolution of ontogenetic changes, population structure and palaeoecology, life‐history strategies, and heterochrony along evolutionary lineages that would otherwise be lost to time. Saurian reptiles (the least‐inclusive clade containing Lepidosauria and Archosauria) have remained an incredibly diverse, numerous, and disparate clade through their ~260‐million‐year history. Because of the great disparity in this group, assessing maturity of saurian reptiles is difficult, fraught with methodological and terminological ambiguity. We compiled a novel database of literature, assembling >900 individual instances of saurian maturity assessment, to examine critically how saurian maturity has been diagnosed. We review the often inexact and inconsistent terminology used in saurian maturity assessment (e.g. ‘juvenile’, ‘mature’) and provide routes for better clarity and cross‐study coherence. We describe the various methods that have been used to assess maturity in every major saurian group, integrating data from both extant and extinct taxa to give a full account of the current state of the field and providing method‐specific pitfalls, best practices, and fruitful directions for future research. We recommend that a new standard subsection, ‘Ontogenetic Assessment’, be added to the Systematic Palaeontology portions of descriptive studies to provide explicit ontogenetic diagnoses with clear criteria. Because the utility of different ontogenetic criteria is highly subclade dependent among saurians, even for widely used methods (e.g. neurocentral suture fusion), we recommend that phylogenetic context, preferably in the form of a phylogenetic bracket, be used to justify the use of a maturity assessment method. Different methods should be used in conjunction as independent lines of evidence when assessing maturity, instead of an ontogenetic diagnosis resting entirely on a single criterion, which is common in the literature. Critically, there is a need for data from extant taxa with well‐represented growth series to be integrated with the fossil record to ground maturity assessments of extinct taxa in well‐constrained, empirically tested methods.

Mass extinctions change global ecosystems, and the end‐Triassic mass extinction was hypothesized to have precipitated the rise of dinosaur dominance, with dinosaurs filling resource zones of eliminated large‐bodied reptilian lineages. This replacement has been explicitly hypothesized for theropod dinosaurs, and the eastern North American theropod footprint record suggests an increase in maximum body size across the Triassic‐Jurassic boundary. Without taking ontogenetic stage in account, the maximum size of the rare large Triassic theropods worldwide supports this hypothesis, with the size of the largest individuals corresponding to the largest Triassic theropod tracks. However, both morphological data and histological examination suggest that known large‐bodied Triassic theropods are represented by immature individuals still growing rapidly at the time of death, indicating that the maximum body size of Triassic theropods was much larger than that a strict reading of the body fossil record would suggest. The size increase recorded in the sediments of eastern North America is not part of a global trend. Instead of a simple ecological replacement of non‐dinosaurian archosaurs by dinosaurs, the rise in theropod dinosaurian ecological dominance was an extended process across the end of the Late Triassic into the Jurassic. Anat Rec, 303:1158–1169, 2020. © 2019 Wiley Periodicals, Inc.

The sacrum - consisting of those vertebrae that articulate with the ilia - is the exclusive skeletal connection between the hindlimbs and axial skeleton in tetrapods. Therefore, the morphology of this portion of the vertebral column plays a major role in the evolution of terrestrial locomotion. Whereas most extant reptiles only possess the two plesiomorphic sacral vertebrae, additional vertebrae have been incorporated into the sacrum multiple times independently among early-diverging archosaurian (crocodylians + birds) clades. Phytosauria was a diverse, abundant, and cosmopolitan clade of archosauriforms throughout the Late Triassic, but postcrania of this clade are rarely described and few species-level taxonomic placements of phytosaurian postcranial material are available, potentially hampering knowledge of morphological disparity in the postcranial skeleton among phytosaurs. Here, we describe the sacrum of Smilosuchus adamanensis, a phytosaur recovered from the Upper Triassic Chinle Formation of Arizona. This sacrum consists of the two primordial sacral vertebrae, but has a vertebra incorporated from the trunk into the sacrum (= a dorsosacral) and is therefore the first Late Triassic phytosaur and one of the first non-archosaurian archosauromorphs to be described with more than two sacral vertebrae. Our interpretation of this element as a dorsosacral is justified by the lateral extent of the dorsosacral ribs, clear surfaces of articulation between the distal ends of the dorsosacral ribs and the first primordial sacral ribs, and the scar on the medial surface of each ilium for articulation with each dorsosacral rib. Additionally, we provide the first detailed description of the vertebral junction formed by two anteriorly projecting flanges on the first primordial sacral ribs and their corresponding facets on the centrum of the dorsosacral. Computed tomographic (CT) imaging reveals that the two primordial sacrals are not co-ossified and that the dorsosacral morphology of this specimen is not the result of obvious pathology. We place this incorporation of a trunk vertebra into the phytosaurian sacrum in a broader evolutionary context, with this shift in vertebral identity occurring at least seven times independently among Triassic archosauriforms, including at least three times in early crocodylian-line archosaurs and at least four times among bird-line archosaurs. Additionally, anteriorly projecting flanges of sacral ribs which articulate with the anterior-adjacent centrum have evolved several times in archosauriforms, and we interpret 'shared' sacral ribs (= a sacral rib that articulates with two adjacent sacral centra more or less equally) present in some archosaurian clades as a more extreme example of this morphology. In extant taxa the highly conserved Hox gene family plays a central role in the patterning of the axial skeleton, especially vertebral identity; therefore, the independent incorporation of a trunk vertebra into the sacrum across multiple archosauriform lineages may suggest a homologous underly...

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Large body sizes among nonavian theropod dinosaurs is a major feature in the evolution of this clade, with theropods reaching greater sizes than any other terrestrial carnivores. However, the early evolution of large body sizes among theropods is obscured by an incomplete fossil record, with the largest Triassic theropods represented by only a few individuals of uncertain ontogenetic stage. Here I describe two neotheropod specimens from the Upper Triassic Bull Canyon Formation of New Mexico and place them in a broader comparative context of early theropod anatomy. These specimens possess morphologies indicative of ontogenetic immaturity (e.g., absence of femoral bone scars, lack of co-ossification between the astragalus and calcaneum), and phylogenetic analyses recover these specimens as early-diverging neotheropods in a polytomy with other early neotheropods at the base of the clade. Ancestral state reconstruction for body size suggests that the ancestral theropod condition was small (~240 mm femur length), but the ancestral neotheropod was larger (~300–340 mm femur length), with coelophysoids experiencing secondary body size reduction, although this is highly dependent on the phylogenetic position of a few key taxa. Theropods evolved large body sizes before the Triassic–Jurassic extinction, as hypothesized in most other ancestral state reconstructions of theropod body sizes, but remained rare relative to smaller theropods until the Jurassic.