Shed teeth from Portezuelo formation at Sierra del Portezuelo reveal a higher diversity of predator theropods during Turonian-Coniacian times in northern Patagonia.

The study of thirty-two shed crowns from the Portezuelo Formation (middle Turonian-late Coniacian) at the Sierra del Portezuelo locality, reveals six distinct tooth morphotypes identified through cladistic, discriminant, and cluster analyses. Two morphotypes were identified as belonging to Megaraptoridae, three to Abelisauridae, one to Abelisauroidea, and one to Alvarezsauridae. Additionally, two of the morphotypes exhibit a combination of dental features typically found in megaraptorid and abelisauridtheropods. These results suggest a greater diversity of theropods in the original ecosystem than previously thought, including the presence of a second morphotype of megaraptorid and alvarezsaurid previously undocumented in this formation. Furthermore, the existence of Morphotype 6 indicates the potential coexistence of medium-sized abelisauroids alongside larger abelisaurids in the same ecosystem. These findings underscore the importance of future expeditions to the Sierra del Portezuelo locality to further our understanding of these previously unknown theropod species.

New information on paleopathologies in non-avian theropod dinosaurs: a case study on South American abelisaurids.

Studies on pathological fossil bones have allowed improving the knowledge of physiology and ecology, and consequently the life history of extinct organisms. Among extinct vertebrates, non-avian dinosaurs have drawn attention in terms of pathological evidence, since a wide array of fossilized lesions and diseases were noticed in these ancient organisms. Here, we evaluate the pathological conditions observed in individuals of different brachyrostran (Theropoda, Abelisauridae) taxa, including Aucasaurus garridoi, Elemgasem nubilus, and Quilmesaurus curriei. For this, we use multiple methodological approaches such as histology and computed tomography, in addition to the macroscopic evaluation. The holotype of Aucasaurus shows several pathognomonic traits of a failure of the vertebral segmentation during development, causing the presence of two fused caudal vertebrae. The occurrence of this condition in Aucasaurus is the first case to be documented so far in non-tetanuran theropods. Regarding the holotype of Elemgasem, the histology of two fused vertebrae shows an intervertebral space between the centra, thus the fusion is limited to the distal rim of the articular surfaces. This pathology is here considered as spondyloarthropathy, the first evidence for a non-tetanuran theropod. The microstructural arrangement of the right tibia of Quilmesaurus shows a marked variation in a portion of the outer cortex, probably due to the presence of the radial fibrolamellar bone tissue. Although similar bone tissue is present in other extinct vertebrates and the cause of its formation is still debated, it could be a response to some kind of pathology. Among non-avian theropods, traumatic injuries are better represented than other maladies (e.g., infection, congenital or metabolic diseases, etc.). These pathologies are recovered mainly among large-sized theropods such as Abelisauridae, Allosauridae, Carcharodontosauridae, and Tyrannosauridae, and distributed principally among axial elements. Statistical tests on the distribution of injuries in these theropod clades show a strong association between taxa-pathologies, body regions-pathologies, and taxa-body regions, suggesting different life styles and behaviours may underlie the frequency of different injuries among theropod taxa.

Developmental strategies underlying gigantism and miniaturization in non-avialan theropod dinosaurs.

In amniotes, the predominant developmental strategy underlying body size evolution is thought to be adjustments to the rate of growth rather than its duration. However, most theoretical and experimental studies supporting this axiom focus on pairwise comparisons and/or lack an explicit phylogenetic framework. We present the first large-scale phylogenetic comparative analysis examining developmental strategies underlying the evolution of body size, focusing on non-avialan theropod dinosaurs. We reconstruct ancestral states of growth rate and body mass in a taxonomically rich dataset, finding that contrary to expectations, changes in the rate and duration of growth played nearly equal roles in the evolution of the vast body size disparity present in non-avialan theropods-and perhaps that of amniotes in general.

The effects of skull flattening on suchian jaw muscle evolution.

Jaw muscles are key features of the vertebrate feeding apparatus. The jaw musculature is housed in the skull whose morphology reflects a compromise between multiple functions, including feeding, housing sensory structures, and defense, and the skull constrains jaw muscle geometry. Thus, jaw muscle anatomy may be suboptimally oriented for the production of bite force. Crocodylians are a group of vertebrates that generate the highest bite forces ever measured with a flat skull suited to their aquatic ambush predatory style. However, basal members of the crocodylian line (e.g., Prestosuchus) were terrestrial predators with plesiomorphically tall skulls, and thus the origin of modern crocodylians involved a substantial reorganization of the feeding apparatus and its jaw muscles. Here, we reconstruct jaw muscles across a phylogenetic range of crocodylians and fossil suchians to investigate the impact of skull flattening on muscle anatomy. We used imaging data to create 3D models of extant and fossil suchians that demonstrate the evolution of the crocodylian skull, using osteological correlates to reconstruct muscle attachment sites. We found that jaw muscle anatomy in early fossil suchians reflected the ancestral archosaur condition but experienced progressive shifts in the lineage leading to Metasuchia. In early fossil suchians, musculus adductor mandibulae posterior and musculus pterygoideus (mPT) were of comparable size, but by Metasuchia, the jaw musculature is dominated by mPT. As predicted, we found that taxa with flatter skulls have less efficient muscle orientations for the production of high bite force. This study highlights the diversity and evolution of jaw muscles in one of the great transformations in vertebrate evolution.

Subaqueous foraging among carnivorous dinosaurs.

Secondary aquatic adaptations evolved independently more than 30 times from terrestrial vertebrate ancestors1,2. For decades, non-avian dinosaurs were believed to be an exception to this pattern. Only a few species have been hypothesized to be partly or predominantly aquatic3-11. However, these hypotheses remain controversial12,13, largely owing to the difficulty of identifying unambiguous anatomical adaptations for aquatic habits in extinct animals. Here we demonstrate that the relationship between bone density and aquatic ecologies across extant amniotes provides a reliable inference of aquatic habits in extinct species. We use this approach to evaluate the distribution of aquatic adaptations among non-avian dinosaurs. We find strong support for aquatic habits in spinosaurids, associated with a marked increase in bone density, which precedes the evolution of more conspicuous anatomical modifications, a pattern also observed in other aquatic reptiles and mammals14-16. Spinosaurids are revealed to be aquatic specialists with surprising ecological disparity, including subaqueous foraging behaviour in Spinosaurus and Baryonyx, and non-diving habits in Suchomimus. Adaptation to aquatic environments appeared in spinosaurids during the Early Cretaceous, following their divergence from other tetanuran theropods during the Early Jurassic17.

On the homology of crocodylian post-dentary bones and their macroevolution throughout Pseudosuchia.

The lower jaw of early tetrapods is composed of several intramembranous ossifications. However, a tendency toward the independent reduction of the number of bones has been observed in the mandible of mammals, lepidosaurs, turtles, crocodiles, and birds. Regarding archosaurs, the coronoid and prearticular bones are interpreted to be lost during the evolution of stem-birds and stem-crocodiles, respectively, but the homology of the post-dentary bones retained in living pseudosuchians remains unclear. Here, we combine paleontological and embryological evidence to explore in detail the homology of the crocodylian post-dentary bones. We study the mandible embryogenesis on a sample of 71 embryos of Caiman and compare this pattern with the mandibular transformations observed across pseudosuchian evolution. In the pre-hatching ontogeny of caimans, at least five intramembranous ossification centers are formed along the margins of the internal mandibular fenestra (perifenestral centers) and, subsequently, merge to form the coronoid (three intramembranous centers), angular (one intramembranous center), and articular (one intramembranous and one chondral center). In the fossil record, an independent prearticular is lost around the base of Mesoeucrocodylia (optimized as reappearing in Thalattosuchia if they are placed within Neosuchia), and the coronoid is apomorphically lost in notosuchians. The integration of embryological and paleontological data indicates that most perifenestral centers are involved in the origin of the prearticular of non-mesoeucrocodylian pseudosuchians. These centers are rearranged during the evolution to contribute to different post-dentary bones in mesoeucrocodylians bolstering the idea that the coronoid and the articular of Crocodylia are not completely homologous to those of other diapsids.

Biomechanical performance of the cranio-mandibular complex of the small notosuchian Araripesuchus gomesii (Notosuchia, Uruguaysuchidae).

Notosuchia is a clade of crocodyliforms that was highly successful and diverse in the Cretaceous of Gondwana. Araripesuchus gomesii is a small notosuchian from the Early Cretaceous of Brazil that belongs to Uruguaysuchidae, one of the subgroups of notosuchians that first radiated, during the Aptian-Albian. Here we present a finite element analysis of A. gomesii based on a model reconstructed from CT scans and performed using published bone properties for crocodiles. The adductor musculature and their respective attachment areas were reconstructed based on Extant Phylogenetic Bracket. Different functional scenarios were tested applying an estimated 158 N bite force: unilateral bite, bilateral bite, pullback, head-shake, and head-twist. The results obtained were compared with those of Alligator mississippiensis, one of its closest living relatives. In the different simulations, the skull and lower jaws of Araripesuchus suffers more stress in the head-shake movement, followed by the unilateral and pullback bites with stress focalized in the premaxillary region. In contrast, the head-twist is the one with smaller stress values. Araripesuchus possess an oreinirostral skull that may provide greater overall resistance in the different scenarios on average, unlike Alligator that has a platyrostral skull with less resistance to dorsoventral mechanical loads. Previous hypotheses that considered A. gomesii as omnivorous coupled with our results, its small size, and likely limited bite force, suggest this taxon probably fed on small prey and other trophic items that could catch and handle entirely with its mouth, such as insects and small vertebrates.

Sauropodomorph evolution across the Triassic-Jurassic boundary: body size, locomotion, and their influence on morphological disparity.

Sauropodomorph dinosaurs were the dominant medium to large-sized herbivores of most Mesozoic continental ecosystems, being characterized by their long necks and reaching a size unparalleled by other terrestrial animals (> 60 tonnes). Our study of morphological disparity across the entire skeleton shows that during the Late Triassic the oldest known sauropodomorphs occupied a small region of morphospace, subsequently diversifying both taxonomically and ecologically, and shifting to a different and broader region of the morphospace. After the Triassic-Jurassic boundary event, there are no substancial changes in sauropodomorph morphospace occupation. Almost all Jurassic sauropodomorph clades stem from ghost lineages that cross the Triassic-Jurassic boundary, indicating that variations after the extinction were more related to changes of pre-existing lineages (massospondylids, non-gravisaurian sauropodiforms) rather than the emergence of distinct clades or body plans. Modifications in the locomotion (bipedal to quadrupedal) and the successive increase in body mass seem to be the main attributes driving sauropodomorph morphospace distribution during the Late Triassic and earliest Jurassic. The extinction of all non-sauropod sauropodomorphs by the Toarcian and the subsequent diversification of gravisaurian sauropods represent a second expansion of the sauropodomorph morphospace, representing the onset of the flourishing of these megaherbivores that subsequently dominated in Middle and Late Jurassic terrestrial assemblages.

Earliest evidence of herd-living and age segregation amongst dinosaurs.

Sauropodomorph dinosaurs dominated the herbivorous niches during the first 40 million years of dinosaur history (Late Triassic-Early Jurassic), yet palaeobiological factors that influenced their evolutionary success are not fully understood. For instance, knowledge on their behaviour is limited, although herding in sauropodomorphs has been well documented in derived sauropods from the Late Jurassic and Cretaceous. Here we report an exceptional fossil occurrence from Patagonia that includes over 100 eggs and skeletal specimens of 80 individuals of the early sauropodomorph Mussaurus patagonicus, ranging from embryos to fully-grown adults, with an Early Jurassic age as determined by high-precision U-Pb zircon geochronology. Most specimens were found in a restricted area and stratigraphic interval, with some articulated skeletons grouped in clusters of individuals of approximately the same age. Our new discoveries indicate the presence of social cohesion throughout life and age-segregation within a herd structure, in addition to colonial nesting behaviour. These findings provide the earliest evidence of complex social behaviour in Dinosauria, predating previous records by at least 40 My. The presence of sociality in different sauropodomorph lineages suggests a possible Triassic origin of this behaviour, which may have influenced their early success as large terrestrial herbivores.

Complex macroevolutionary dynamics underly the evolution of the crocodyliform skull.

All modern crocodyliforms (alligators, crocodiles and the gharial) are semi-aquatic generalist carnivores that are relatively similar in cranial form and function. However, this homogeneity represents just a fraction of the variation that once existed in the clade, which includes extinct herbivorous and marine forms with divergent skull structure and function. Here, we use high-dimensional three-dimensional geometric morphometrics to quantify whole-skull morphology across modern and fossil crocodyliforms to untangle the factors that shaped the macroevolutionary history and relatively low phenotypic variation of this clade through time. Evolutionary modelling demonstrates that the pace of crocodyliform cranial evolution is initially high, particularly in the extinct Notosuchia, but slows near the base of Neosuchia, with a late burst of rapid evolution in crown-group crocodiles. Surprisingly, modern crocodiles, especially Australian, southeast Asian, Indo-Pacific species, have high rates of evolution, despite exhibiting low variation. Thus, extant lineages are not in evolutionary stasis but rather have rapidly fluctuated within a limited region of morphospace, resulting in significant convergence. The structures related to jaw closing and bite force production (e.g. pterygoid flange and quadrate) are highly variable, reinforcing the importance of function in driving phenotypic variation. Together, these findings illustrate that the apparent conservativeness of crocodyliform skulls betrays unappreciated complexity in their macroevolutionary dynamics.

Unexpected larger distribution of paleogene stem-rollers (AVES, CORACII): new evidence from the Eocene of Patagonia, Argentina.

Here we present the first record of a stem-Coracii outside the Holarctic region, found in the early Eocene of Patagonia at the Laguna del Hunco locality. Ueekenkcoracias tambussiae gen. et sp. nov. consists of an incomplete right hind limb that presents the following combination of characters, characteristic of Coracii: relatively short and stout tibiotarsus, poorly developed crista cnemialis cranialis, short and wide tarsometatarsus, with the tuberositas m. tibialis cranialis located medially on the shaft, and curved and stout ungual phalanges. Although the presence of a rounded and conspicuous foramen vasculare distale and the trochlea metatarsi II strongly deflected medially resemble Primobucconidae, a fossil group only found in the Eocene of Europe and North America, our phylogenetic analysis indicates the new taxon is the basalmost known Coracii. The unexpected presence of a stem-Coracii in the Eocene of South America indicates that this clade had a more widespread distribution than previously hypothesized, already extending into the Southern Hemisphere by the early Eocene. Ueekenkcoracias tambussiae represents new evidence of the increasing diversity of stem lineages of birds in the Eocene. The new material provides novel morphological data for understanding the evolutionary origin and radiation of rollers and important data for estimates of the divergence time of the group.

The first dinosaur egg was soft.

Calcified eggshells protect developing embryos against environmental stress and contribute to reproductive success1. As modern crocodilians and birds lay hard-shelled eggs, this eggshell type has been inferred for non-avian dinosaurs. Known dinosaur eggshells are characterized by an innermost membrane, an overlying protein matrix containing calcite, and an outermost waxy cuticle2-7. The calcitic eggshell consists of one or more ultrastructural layers that differ markedly among the three major dinosaur clades, as do the configurations of respiratory pores. So far, only hadrosaurid, a few sauropodomorph and tetanuran eggshells have been discovered; the paucity of the fossil record and the lack of intermediate eggshell types challenge efforts to homologize eggshell structures across all dinosaurs8-18. Here we present mineralogical, organochemical and ultrastructural evidence for an originally non-biomineralized, soft-shelled nature of exceptionally preserved ornithischian Protoceratops and basal sauropodomorph Mussaurus eggs. Statistical evaluation of in situ Raman spectra obtained for a representative set of hard- and soft-shelled, fossil and extant diapsid eggshells clusters the originally organic but secondarily phosphatized Protoceratops and the organic Mussaurus eggshells with soft, non-biomineralized eggshells. Histology corroborates the organic composition of these soft-shelled dinosaur eggs, revealing a stratified arrangement resembling turtle soft eggshell. Through an ancestral-state reconstruction of composition and ultrastructure, we compare eggshells from Protoceratops and Mussaurus with those from other diapsids, revealing that the first dinosaur egg was soft-shelled. The calcified, hard-shelled dinosaur egg evolved independently at least three times throughout the Mesozoic era, explaining the bias towards eggshells of derived dinosaurs in the fossil record.

Braincase anatomy of Almadasuchus figarii (Archosauria, Crocodylomorpha) and a review of the cranial pneumaticity in the origins of Crocodylomorpha.

Almadasuchus figarii is a basal crocodylomorph recovered from the Upper Jurassic levels of the Cañadón Calcáreo Formation (Oxfordian-Tithonian) of Chubut, Argentina. This taxon is represented by cranial remains, which consist of partial snout and palatal remains; an excellently preserved posterior region of the skull; and isolated postcranial remains. The skull of the only specimen of the monotypic Almadasuchus was restudied using high-resolution computed micro tomography. Almadasuchus has an apomorphic condition in its skull shared with the closest relatives of crocodyliforms (i.e. hallopodids) where the quadrates are sutured to the laterosphenoids and the otoccipital contacts the quadrate posterolaterally, reorganizing the exit of several cranial nerves (e.g. vagus foramen) and the entry of blood vessels (e.g. internal carotids) on the occipital surface of the skull. The endocast is tubular, as previously reported in thalattosuchians, but has a marked posterior step, and a strongly projected floccular recess as in other basal crocodylomorphs. Internally, the skull of Almadasuchus is heavily pneumatized, where different air cavities invade the bones of the suspensorium and braincase, both on its dorsal or ventral parts. Almadasuchus has a large basioccipital recess, which is formed by cavities that excavate the basioccipital and the posterior surface of the basisphenoid, and unlike other crocodylomorphs is connected with the basisphenoid pneumatizations. Ventral to the otic capsule, a pneumatic cavity surrounded by the otoccipital and basisphenoid is identified as the rhomboidal recess. The quadrate of Almadasuchus is highly pneumatized, being completely hollow, and the dorsal pneumatizations of the braincase are formed by the mastoid and facial antra, and a laterosphenoid cavity (trigeminal diverticulum). To better understand the origins of pneumatic features in living crocodylomorphs we studied cranial pneumaticity in the basal members of Crocodylomorpha and found that: (a) prootic pneumaticity may be a synapomorphy for the whole clade; (b) basisphenoid pneumaticity (pre-, postcarotid and rostral recesses) is a derived feature among basal crocodylomorphs; (c) quadrate pneumatization is acquired later in the history of the group; and (d) the rhomboidal sinus is a shared derived trait of hallopodids and crocodyliforms. The marine thallatosuchians exhibit a reduction of the pneumaticity of the braincase and this reduction is evaluated considering the two phylogenetic positions proposed for the clade.

Probable basal allosauroid from the early Middle Jurassic Cañadón Asfalto Formation of Argentina highlights phylogenetic uncertainty in tetanuran theropod dinosaurs.

Tetanurae, the most successful clade of theropod dinosaurs, including modern birds, split into three major clades early in their evolutionary history: Megalosauroidea, Coelurosauria, and Allosauroidea. The oldest tetanurans occur in the earliest Middle Jurassic, but the early fossil record of the clade is still poor. Here we report one of the oldest known and most complete pre-Late Jurassic tetanuran, the probable allosauroid Asfaltovenator vialidadi gen. et sp. nov., which has an unusual character combination, uniting features currently considered to be apomorphic of different tetanuran lineages. A phylogenetic analysis resulted in a monophyletic Carnosauria (Allosauroidea + Megalosauroidea), and the inclusion of the new taxon significantly changes topology within carnosaurs. The analysis shows concentrated homoplasy in proximal nodes at the base of Tetanurae, and a temporal peak at the Pliensbachian-Toarcian extinction event, recently identified as a potential driver of tetanuran radiation. These results highlight the complex morphological evolution in the early radiation of tetanuran theropods, in which convergences and parallelisms were extremely common. This pattern seems to be a common feature in rapid radiation events of major clades of vertebrates and might explain the common difficulties to unravel phylogenetic relationships of important lineages at the base of major clades.

Ontogenetic changes in the body plan of the sauropodomorph dinosaur Mussaurus patagonicus reveal shifts of locomotor stance during growth.

Ontogenetic information is crucial to understand life histories and represents a true challenge in dinosaurs due to the scarcity of growth series available. Mussaurus patagonicus was a sauropodomorph dinosaur close to the origin of Sauropoda known from hatchling, juvenile and mature specimens, providing a sufficiently complete ontogenetic series to reconstruct general patterns of ontogeny. Here, in order to quantify how body shape and its relationship with locomotor stance (quadruped/biped) changed in ontogeny, hatchling, juvenile (~1 year old) and adult (8+ years old) individuals were studied using digital models. Our results show that Mussaurus rapidly grew from about 60 g at hatching to ~7 kg at one year old, reaching >1000 kg at adulthood. During this time, the body's centre of mass moved from a position in the mid-thorax to a more caudal position nearer to the pelvis. We infer that these changes of body shape and centre of mass reflect a shift from quadrupedalism to bipedalism occurred early in ontogeny in Mussaurus. Our study indicates that relative development of the tail and neck was more influential in determining the locomotor stance in Sauropodomorpha during ontogeny, challenging previous studies, which have emphasized the influence of hindlimb vs. forelimb lengths on sauropodomorph stance.

Evolutionary Integration and Modularity in the Archosaur Cranium.

Complex structures, like the vertebrate skull, are composed of numerous elements or traits that must develop and evolve in a coordinated manner to achieve multiple functions. The strength of association among phenotypic traits (i.e., integration), and their organization into highly-correlated, semi-independent subunits termed modules, is a result of the pleiotropic and genetic correlations that generate traits. As such, patterns of integration and modularity are thought to be key factors constraining or facilitating the evolution of phenotypic disparity by influencing the patterns of variation upon which selection can act. It is often hypothesized that selection can reshape patterns of integration, parceling single structures into multiple modules or merging ancestrally semi-independent traits into a strongly correlated unit. However, evolutionary shifts in patterns of trait integration are seldom assessed in a unified quantitative framework. Here, we quantify patterns of evolutionary integration among regions of the archosaur skull to investigate whether patterns of cranial integration are conserved or variable across this diverse group. Using high-dimensional geometric morphometric data from 3D surface scans and computed tomography scans of modern birds (n = 352), fossil non-avian dinosaurs (n = 27), and modern and fossil mesoeucrocodylians (n = 38), we demonstrate that some aspects of cranial integration are conserved across these taxonomic groups, despite their major differences in cranial form, function, and development. All three groups are highly modular and consistently exhibit high integration within the occipital region. However, there are also substantial divergences in correlation patterns. Birds uniquely exhibit high correlation between the pterygoid and quadrate, components of the cranial kinesis apparatus, whereas the non-avian dinosaur quadrate is more closely associated with the jugal and quadratojugal. Mesoeucrocodylians exhibit a slightly more integrated facial skeleton overall than the other grades. Overall, patterns of trait integration are shown to be stable among archosaurs, which is surprising given the cranial diversity exhibited by the clade. At the same time, evolutionary innovations such as cranial kinesis that reorganize the structure and function of complex traits can result in modifications of trait correlations and modularity.

Morphological Data Sets Fit a Common Mechanism Much More Poorly than DNA Sequences and Call Into Question the Mkv Model.

The Mkv evolutionary model, based on minor modifications to models of molecular evolution, is being increasingly used to infer phylogenies from discrete morphological data, often producing different results from parsimony. The critical difference between Mkv and parsimony is the assumption of a "common mechanism" in the Mkv model, with branch lengths determining that probability of change for all characters increases or decreases at the same tree branches by the same exponential factor. We evaluate whether the assumption of a common mechanism applies to morphology, by testing the implicit prediction that branch lengths calculated from different subsets of characters will be significantly correlated. Our analysis shows that DNA (38 data sets tested) is often compatible with a common mechanism, but morphology (86 data sets tested) generally is not, showing very disparate branch lengths for different character partitions. The low levels of branch length correlation demonstrated for morphology (fitting models without a common mechanism) suggest that the Mkv model is too unrealistic and inadequate for the analysis of most morphological data sets. [Bayesian analysis; Mkv model; morphological data; phylogenetics.].

The Long Bone Histology of the Sauropodomorph, Antetonitrus ingenipes.

This analysis of the long bone microstructure of Antetonitrus ingenipes fills a crucial gap in our understanding of the growth dynamics of sauropodomorph dinosaurs. The bone histology of basal Sauropodomorpha are often characterized by zonal tissue, and contrasts with that of more derived sauropod taxa which show a shift toward the deposition of uninterrupted fibrolamellar bone (with lines of growth being either absent or only present in the outer circumferential layer). In Antetonitrus, growth patterns in the youngest individuals exhibit uninterrupted fibrolamellar bone without any growth marks. Sub-adult individuals, also exhibit highly vascularized fibrolamellar bone throughout the cortex, as in more derived Sauropods and Mussaurus, but growth lines occur intermittently (although not regularly) throughout the cortex as in Lessemsaurus. This indicates that Antetonitrus does not exhibit the growth dynamics previously considered characteristic of Sauropoda. Despite this, the largest (and possibly the oldest femur, NMQR 1705/163) does show an incipient external fundamental system (EFS). Our findings further suggest that growth marks are decoupled from bone size, which indicates a level of developmental plasticity in this taxon. Modulations or textural shifts in the pattern of vascular channel arrangements throughout the fibrolamellar bone in the cortex may be related to periods of resource limitations, although the lack of consistency of these modulations suggest that it is unlikely due to seasonal fluctuations. Localized bands of radial fibrolamellar bone, followed by resumption of normal growth in two samples are interpreted as evidence of a disease infliction, and subsequent recovery thereof. Anat Rec, 301:1506-1518, 2018. © 2018 Wiley Periodicals, Inc.

An early trend towards gigantism in Triassic sauropodomorph dinosaurs.

Dinosaurs dominated the terrestrial ecosystems for more than 140 Myr during the Mesozoic era, and among them were sauropodomorphs, the largest land animals recorded in the history of life. Early sauropodomorphs were small bipeds, and it was long believed that acquisition of giant body size in this clade (over 10 tonnes) occurred during the Jurassic and was linked to numerous skeletal modifications present in Eusauropoda. Although the origin of gigantism in sauropodomorphs was a pivotal stage in the history of dinosaurs, an incomplete fossil record obscures details of this crucial evolutionary change. Here, we describe a new sauropodomorph from the Late Triassic of Argentina nested within a clade of other non-eusauropods from southwest Pangaea. Members of this clade attained large body size while maintaining a plesiomorphic cyclical growth pattern, displaying many features of the body plan of basal sauropodomorphs and lacking most anatomical traits previously regarded as adaptations to gigantism. This novel strategy highlights a highly accelerated growth rate, an improved avian-style respiratory system, and modifications of the vertebral epaxial musculature and hindlimbs as critical to the evolution of gigantism. This reveals that the first pulse towards gigantism in dinosaurs occurred over 30 Myr before the appearance of the first eusauropods.