Neurocranial osteology and neuroanatomy of a Late Cretaceous Titanosaurian sauropod from Spain (Ampelosaurus sp.)

Titanosaurians were a flourishing group of sauropod dinosaurs during Cretaceous times. Fossils of titanosaurians have been found on all continents and their remains are abundant in a number of Late Cretaceous sites. Nonetheless, the cranial anatomy of titanosaurians is still very poorly known. The Spanish latest Cretaceous locality of >Lo Hueco> yielded a relatively well preserved, titanosaurian braincase, which shares a number of phylogenetically restricted characters with Ampelosaurus atacis from France such as a flat occipital region. However, it appears to differ from A. atacis in some traits such as the greater degree of dorsoventral compression and the presence of proatlas facets. The specimen is, therefore, provisionally identified as Ampelosaurus sp. It was CT scanned, and 3D renderings of the cranial endocast and inner-ear system were generated. Our investigation highlights that, although titanosaurs were derived sauropods with a successful evolutionary history, they present a remarkably modest level of paleoneurological organization. Compared with the condition in the basal titanosauriform Giraffatitan brancai, the labyrinth of Ampelosaurus sp. shows a reduced morphology. The latter feature is possibly related to a restricted range of head-turning movements. © 2013 Knoll et al.This is a contribution to the research project CGL2009-12143 (Ministerio de Economı´a y Competitividad, Madrid), of which FK, who is currently supported by the Ramo´n y Cajal Program, is Principal Investigator.Peer Reviewe

Mechanical analysis of feeding behavior in the extinct "Terror Bird" <i>Andalgalornis steulleti</i> (Gruiformes: phorusrhacidae)

The South American phorusrhacid bird radiation comprised at least 18 species of small to gigantic terrestrial predators for which there are no close modern analogs. Here we perform functional analyses of the skull of the medium-sized (,40 kg) patagornithine phorusrhacid Andalgalornis steulleti (upper Miocene-lower Pliocene, Andalgalá Formation, Catamarca, Argentina) to assess its mechanical performance in a comparative context. Based on computed tomographic (CT) scanning and morphological analysis, the skull of Andalgalornis steulleti is interpreted as showing features reflecting loss of intracranial immobility. Discrete anatomical attributes permitting such cranial kinesis are widespread phorusrhacids outgroups, but this is the first clear evidence of loss of cranial kinesis in a gruiform bird and may be among the best documented cases among all birds. This apomorphic loss is interpreted as an adaptation for enhanced craniofacial rigidity, particularly with regard to sagittal loading. We apply a Finite Element approach to a three-dimensional (3D) model of the skull. Based on regression analysis we estimate the bite force of Andalgalornis at the bill tip to be 133 N. Relative to results obtained from Finite Element Analysis of one of its closest living relatives (seriema) and a large predatory bird (eagle), the phorusrhacid's skull shows relatively high stress under lateral loadings, but low stress where force is applied dorsoventrally (sagittally) and in ''pullback'' simulations. Given the relative weakness of the skull mediolaterally, it seems unlikely that Andalgalornis engaged in potentially risky behaviors that involved subduing large, struggling prey with its beak. We suggest that it either consumed smaller prey that could be killed and consumed more safely (e.g., swallowed whole) or that it used multiple well-targeted sagittal strikes with the beak in a repetitive attack-and-retreat strategy.Facultad de Ciencias Naturales y Muse

Cranial osteology of the ankylosaurian dinosaur formerly known as Minmi sp (Ornithischia: Thyreophora) from the Lower Cretaceous Allaru Mudstone of Richmond, Queensland, Australia

Minmi is the only known genus of ankylosaurian dinosaur from Australia. Seven specimens are known, all from the Lower Cretaceous of Queensland. Only two of these have been described in any detail: the holotype specimenMinmi paravertebra fromthe Bungil Formation near Roma, and a near complete skeleton fromthe Allaru Mudstone onMarathon Station near Richmond, preliminarily referred to a possible new species of Minmi. The Marathon specimen represents one of the world's most complete ankylosaurian skeletons and the best-preserved dinosaurian fossil from eastern Gondwana. Moreover, among ankylosaurians, its skull is one of only a few in which the majority of sutures have not been obliterated by dermal ossifications or surface remodelling. Recent preparation of theMarathon specimen has revealed new details of the palate and narial regions, permitting a comprehensive description and thus providing new insights cranial osteology of a basal ankylosaurian. The skull has also undergone computed tomography, digital segmentation and 3D computer visualisation enabling the reconstruction of its nasal cavity and endocranium. The airways of the Marathon specimen are more complicated than non-ankylosaurian dinosaurs but less so than derived ankylosaurians. The cranial (brain) endocast is superficially similar to those of other ankylosaurians but is strongly divergent in many important respects. The inner ear is extremely large and unlike that of any dinosaur yet known. Based on a high number of diagnostic differences between the skull of theMarathon specimen and other ankylosaurians, we consider it prudent to assign this specimen to a new genus and species of ankylosaurian. Kunbarrasaurus ieversi gen. et sp. nov. represents the second genus of ankylosaurian from Australia and is characterised by an unusual melange of both primitive and derived characters, shedding new light on the evolution of the ankylosaurian skull

Cartilaginous Epiphyses in Extant Archosaurs and Their Implications for Reconstructing Limb Function in Dinosaurs

Extinct archosaurs, including many non-avian dinosaurs, exhibit relatively simply shaped condylar regions in their appendicular bones, suggesting potentially large amounts of unpreserved epiphyseal (articular) cartilage. This “lost anatomy” is often underappreciated such that the ends of bones are typically considered to be the joint surfaces, potentially having a major impact on functional interpretation. Extant alligators and birds were used to establish an objective basis for inferences about cartilaginous articular structures in such extinct archosaur clades as non-avian dinosaurs. Limb elements of alligators, ostriches, and other birds were dissected, disarticulated, and defleshed. Lengths and condylar shapes of elements with intact epiphyses were measured. Limbs were subsequently completely skeletonized and the measurements repeated. Removal of cartilaginous condylar regions resulted in statistically significant changes in element length and condylar breadth. Moreover, there was marked loss of those cartilaginous structures responsible for joint architecture and congruence. Compared to alligators, birds showed less dramatic, but still significant changes. Condylar morphologies of dinosaur limb bones suggest that most non-coelurosaurian clades possessed large cartilaginous epiphyses that relied on the maintenance of vascular channels that are otherwise eliminated early in ontogeny in smaller-bodied tetrapods. A sensitivity analysis using cartilage correction factors (CCFs) obtained from extant taxa indicates that whereas the presence of cartilaginous epiphyses only moderately increases estimates of dinosaur height and speed, it has important implications for our ability to infer joint morphology, posture, and the complicated functional movements in the limbs of many extinct archosaurs. Evidence suggests that the sizes of sauropod epiphyseal cartilages surpassed those of alligators, which account for at least 10% of hindlimb length. These data suggest that large cartilaginous epiphyses were widely distributed among non-avian archosaurs and must be considered when making inferences about locomotor functional morphology in fossil taxa

The braincase and inner ear of ‘Metriorhynchus’ cf. ‘M.’ brachyrhynchus – implications for aquatic sensory adaptations in crocodylomorphs

During their long evolutionary history crocodylomorphs achieved a great diversity of body sizes, ecomorphotypes and inferred feeding ecologies. One unique group of crocodylomorphs are the thalattosuchians, which lived during the Jurassic and Cretaceous (ca. 191–125 Ma). They transitioned from shallow marine species, like teleosauroids, into fully pelagic forms with paddle shaped limbs and a vertically orientated tail fluke, the metriorhynchids. The osteological adaptations that allowed metriorhynchids to live in the water are generally well understood, but less is known about their neurosensory and endocranial systems, such as the brain, inner ears, sinuses and cranial nerves and how they relate to their aquatic lifestyle. Based on micro-computed tomography (μCT) data and three-dimensional models, we here describe the braincase and endocranial anatomy of a fully marine metriorhynchid, ‘Metriorhynchus’ cf. ‘M.’ brachyrhynchus (NHMUK PV OR 32617). We found several neuroanatomical features that likely helped this species function in its marine environment. These include a unique flexure in the brain endocast not seen in other thalattosuchians. Other features that have previously been seen in thalattosuchians include enlarged cerebral hemispheres, a hypertrophied venous sinus system, enlarged internal carotid arteries and foramina, and closed/absent lateral pharyngotympanic foramina. The specimen also possesses a pelagic metriorhynchid bony labyrinth morphology, with a compact and dorsoventrally short shape, thick semicircular canals, an enlarged vestibule and potentially a short cochlear duct. A review of character distribution confirms that some of these features evolved at the base of Thalattosuchia in semiaquatic species, long before metriorhynchids became pelagic, suggesting that endocranial anatomy helped allow metriorhynchoids colonize the ocean realm.Fil: Schwab, Julia A.. University of Edinburgh; Reino UnidoFil: Young, Mark T.. University of Edinburgh; Reino UnidoFil: Herrera, Laura Yanina. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Paleontología Vertebrados; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata; ArgentinaFil: Witmer, Lawrence. Ohio University; Estados UnidosFil: Walsh, Stig A.. University of Southampton; Reino UnidoFil: Katsamensis, Orestis. Faculty Of Engineering And Physical Sciences; Reino UnidoFil: Brusatte, Stephen L.. University of Edinburgh; Reino Unid

Edentulism, beaks and biomechanical innovations in the early evolution of theropod dinosaurs

Maniraptoriformes, the speciose group of derived theropod dinosaurs that ultimately gave rise to modern birds, display a diverse and remarkable suite of skeletal adaptations. Apart from the evolution of flight, a large-scale change in dietary behavior appears to have been one of the main triggers for specializations in the bauplan of these derived theropods. Among the different skeletal specializations, partial or even complete edentulism and the development of keratinous beaks form a recurring and persistent trend in from the evolution of derived nonavian dinosaurs. Therizinosauria is an enigmatic maniraptoriform clade, whose members display these and other osteological characters thought to be correlated with the shift from carnivory to herbivory. This makes therizinosaurians prime candidates to assess the functional significance of these morphological characters. Based on a highly detailed biomechanical model of Erlikosaurus andrewsi, a therizinosaurid from the Upper Cretaceous of Mongolia, different morphological configurations incorporating soft-tissue structures, such as a keratinous rhamphotheca, are evaluated for their biomechanical performance. Our results indicate that the development of beaks and the presence of a keratinous rhamphotheca would have helped to dissipate stress and strain, making the rostral part of the skull less susceptible to bending and displacement, and this benefit may extend to other vertebrate clades that possess rhamphothecae. Keratinous beaks, paralleled by edentulism, thus represent an evolutionary innovation developed early in derived theropods to enhance cranial stability, distinct to postulated mass-saving benefits associated with the origin of flight

Ontogenetic variation in the crocodylian vestibular system

Crocodylians today live in tropical to subtropical environments, occupying mostly shallow waters. Their body size changes drastically during ontogeny, as do their skull dimensions and bite forces, which are associated with changes in prey preferences. Endocranial neurosensory structures have also shown to change ontogenetically, but less is known about the vestibular system of the inner ear. Here we use 30 high-resolution computed tomography (CT) scans and three-dimensional geometric morphometrics to investigate the size and shape changes of crocodylian endosseous labyrinths throughout ontogeny, across four stages (hatchling, juvenile, subadult and adult). We find two major patterns of ontogenetic change. First, the labyrinth increases in size during ontogeny, with negative allometry in relation to skull size. Second, labyrinth shape changes significantly, with hatchlings having shorter semicircular canal radii, with thicker diameters and an overall dorsoventrally shorter labyrinth than those of more mature individuals. We argue that the modification of the labyrinth during crocodylian ontogeny is related to constraints imposed by skull growth, due to fundamental changes in the crocodylian braincase during ontogeny (e.g. verticalisation of the basicranium), rather than changes in locomotion, diet, or other biological functions or behaviours

‘Ear stones’ in crocodylians: a cross-species comparative and ontogenetic survey of otolith structures

The vestibular system of the inner ear is a crucial sensory organ, involved in the sensation of balance and equilibrium. It consists of three semicircular canals that sense angular rotations of the head and the vestibule that detects linear acceleration and gravity. The vestibule often contains structures, known as the otoliths or ‘ear stones’. Otoliths are present in many vertebrates and are particularly well known from the fossil record of fish, but surprisingly have not been described in detail in most tetrapods, living or extinct. Here, we present for the first time a survey of the otoliths of a broad sample of extant crocodylian species, based on computed tomography scans. We find that otoliths are present in numerous crocodylian species of different growth stages, and they continue to increase in size during ontogeny, with positive allometry compared to skull length. Our results confirm that otoliths are a common component of the crocodylian vestibular system, and suggest they play an important role in sensory detection. Otoliths are likely common, but overlooked, constituents of the inner ear in tetrapods, and a broader study of their size, shape and distribution promises insight into sensory abilities

Cephalic salt gland evolution in Mesozoic pelagic crocodylomorphs

Secondarily marine tetrapod lineages have independently evolved osmoregulatory adaptations for life in salt water but inferring physiological changes in extinct marine tetrapods is difficult. The Mesozoic crocodylomorph clade Thalattosuchia is unique in having both direct evidence from natural endocasts and several proposed osteological correlates for salt exocrine glands. Here, we investigate salt gland evolution in thalattosuchians by creating endocranial reconstructions from CT scans of eight taxa (one basal thalattosuchian, one teleosauroid, two basal metriorhynchoids and four metriorhynchids) and four outgroups (three extant crocodylians and the basal crocodyliform Protosuchus) to identify salt gland osteological correlates. All metriorhynchoids show dorsolateral nasal cavity expansions corresponding to the location of nasal salt glands in natural casts, but smaller expansions in teleosauroids correspond more with the cartilaginous nasal capsule. The different sizes of these expansions suggest the following evolutionary sequence: (1) plesiomorphically small glands present in semi-aquatic teleosauroids draining through the nasal vestibule; (2) moderately sized glands in the basalmost metriorhynchoid Pelagosaurus; and (3) hypertrophied glands in the clade comprising Eoneustes and metriorhynchids, with a pre-orbital fenestra providing a novel exit for salt drainage. The large gland size inferred from basal metriorhynchoids indicates advanced osmoregulation occurred while metriorhynchoids were semi-aquatic. This pattern does not precisely fit into current models of physiological evolution in marine tetrapods and suggests a unique sequence of changes as thalattosuchians transitioned from land to sea.Fil: Cowgill, T.. University of Edinburgh; Reino UnidoFil: Young, M.. University of Edinburgh; Reino UnidoFil: Schwab, J.. University of Edinburgh; Reino UnidoFil: Walsh, S.. National Museum Of Scotland; Reino UnidoFil: Witmer, Lawrence. Ohio University; Estados UnidosFil: Herrera, Laura Yanina. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Paleontología Vertebrados; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata; ArgentinaFil: Dollman, K.. University of the Witwatersrand; SudáfricaFil: Turner, A. H.. State University of New York. Stony Brook University; Estados UnidosFil: Brusatte, S.. University of Edinburgh; Reino Unid

Cephalic salt gland evolution in Mesozoic pelagic crocodylomorphs

Cowgill, Thomas, Young Fls, Mark T, Schwab, Julia A, Walsh, Stig, Witmer, Lawrence M, Herrera, Yanina, Dollman, Kathleen N, Turner, Alan H, Brusatte, Stephen L (2023): Cephalic salt gland evolution in Mesozoic pelagic crocodylomorphs. Zoological Journal of the Linnean Society 197 (3): 812-835, DOI: 10.1093/zoolinnean/zlac027, URL: https://academic.oup.com/zoolinnean/article/197/3/812/659333