Oldest preserved umbilical scar reveals dinosaurs had ‘belly buttons’

Background: In egg-laying amniotes, the developing embryo is tethered to a number of the extraembryonic membranes including the yolk sac and allantois that deliver oxygen and nutrients and remove metabolic waste products throughout embryonic development. Prior to, or soon after hatching, these membranes detach from the animal leaving a temporary or permanent umbilical scar (umbilicus) equivalent to the navel or ‘belly button’ in some placental mammals, including humans. Although ubiquitous in modern mammals and reptiles (including birds), at least early in their ontogeny, the umbilicus has not been identified in any pre-Cenozoic amniote. Results: We report the oldest preserved umbilicus in a fossil amniote from a ~130-million-year-old early-branching ceratopsian dinosaur, Psittacosaurus. Under laser-stimulated fluorescence (LSF), the umbilicus is revealed as an elongate midline structure delimited by a row of paired scales on the abdomen. The relatively late ontogenetic stage (close to sexual maturity) estimated for the individual indicates that the umbilicus was probably retained throughout life. Conclusions: Unlike most extant reptiles and birds that lose this scar within days to weeks after hatching, the umbilicus of Psittacosaurus persisted at least until sexual maturity, similar to some lizards and crocodylians with which it shares the closest morphological resemblance. This discovery is the oldest record of an amniote umbilicus and the first in a non-avian dinosaur. However, given the variability of this structure in extant reptilian analogues, a persistent umbilical scar may not have been present in all non-avian dinosaurs.Fil: Bell, Phil R.. University Of New England Australia; AustraliaFil: Hendrickx, Christophe Marie Fabian. Fundación Miguel Lillo; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico - Tucumán. Unidad Ejecutora Lillo; ArgentinaFil: Pittman, Michael. Colegio Universitario de Londres; Reino UnidoFil: Kaye, Thomas G.. Foundation For Scientific Advancement; Estados Unido

The exquisitely preserved integument of Psittacosaurus and the scaly skin of ceratopsian dinosaurs

The Frankfurt specimen of the early-branching ceratopsian dinosaur Psittacosaurus is remarkable for the exquisite preservation of squamous (scaly) skin and other soft tissues that cover almost its entire body. New observations under Laser-Stimulated Fluorescence (LSF) reveal the complexity of the squamous skin of Psittacosaurus, including several unique features and details of newly detected and previously-described integumentary structures. Variations in the scaly skin are found to be strongly regionalized in Psittacosaurus. For example, feature scales consist of truncated cone-shaped scales on the shoulder, but form a longitudinal row of quadrangular scales on the tail. Re-examined through LSF, the cloaca of Psittacosaurus has a longitudinal opening, or vent; a condition that it shares only with crocodylians. This implies that the cloaca may have had crocodylian-like internal anatomy, including a single, ventrally-positioned copulatory organ. Combined with these new integumentary data, a comprehensive review of integument in ceratopsian dinosaurs reveals that scalation was generally conservative in ceratopsians and typically consisted of large subcircular-to-polygonal feature scales surrounded by a network of smaller non-overlapping polygonal basement scales. This study highlights the importance of combining exceptional specimens with modern imaging techniques, which are helping to redefine the perceived complexity of squamation in ceratopsians and other dinosaurs.Fil: Bell, Phil R.. University Of New England; AustraliaFil: Hendrickx, Christophe Marie Fabian. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico - Tucumán. Unidad Ejecutora Lillo; Argentina. Fundación Miguel Lillo; ArgentinaFil: Pittman, Michael. Chinese University Of Hong Kong; Hong Kong. Foundation For Scientific Advancement; Estados UnidosFil: Kaye, Thomas G.. Foundation For Scientific Advancement; Estados UnidosFil: Mayr, Gerald. Senckenberg Forschungsinstitut Und Naturmuseum; Alemani

Dental anatomy of the apex predator Sinraptor Dongi (Theropoda: Allosauroidea) from the late Jurassic of China

The dental morphology of the holotype of the theropod Sinraptor dongi from the Jurassic Shishugou Formation of China is comprehensively described. We highlight a combination of dental features that appear to be restricted to Sinraptor: (i) crowns with denticulated mesial and distal carinae extending from the root and an irregular surface texture on the enamel; (ii) a D- to salinon-shaped cross-sectional outline at the crown base in mesialmost teeth; (iii) mesial crowns with mesial carinae spiraling mesiolingually and lingually positioned longitudinal groove adjacent to the mesial carina; and (iv) particularly labiolingually compressed lateral teeth with weakly labially deflected distal carinae, flat to concave basocentral surfaces of the labial margins of the crowns, and horizontally elongated distal denticles showing short to well-developed interdenticular sulci. Using cladistic, multivariate, discriminant, and cluster analyses, we demonstrate that the dentition of Sinraptor is relatively similar to that of ceratosaurids, megalosauroids, and other allosauroids and is particularly close to that of Allosaurus. The dental anatomy of Sinraptor and Allosaurus, which differs mainly in the labiolingual compression of the lateral crowns and in the number of premaxillary teeth, shows adaptations towards a predatory lifestyle, including premaxillary teeth capable of enduring tooth-tobone contact and crowns with widely separated mesial and distal carinae capable of inflicting widely open wounds.Fil: Hendrickx, Christophe Marie Fabian. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico - Tucumán. Unidad Ejecutora Lillo; ArgentinaFil: Stiegler, Josef. The George Washington University; Estados UnidosFil: Currie, Philip J.. University of Alberta; CanadáFil: Han, Fenglu. University of Geoscience; ChinaFil: Xu, Xing. Chinese Academy of Sciences; República de ChinaFil: Choiniere, Jonah N.. University of the Witwatersrand; SudáfricaFil: Wu, Xiao Chung. Canadian Museum of Nature; Canad

Morphology and distribution of scales, dermal ossifications, and other non-feather integumentary structures in non-avialan theropod dinosaurs

Modern birds are typified by the presence of feathers, complex evolutionary innovations that were already widespread in the group of theropod dinosaurs (Maniraptoriformes) that include crown Aves. Squamous or scaly reptilian-like skin is, however, considered the plesiomorphic condition for theropods and dinosaurs more broadly. Here, we review the morphology and distribution of non-feathered integumentary structures in non-avialan theropods, covering squamous skin and naked skin as well as dermal ossifications. The integumentary record of non-averostran theropods is limited to tracks, which ubiquitously show a covering of tiny reticulate scales on the plantar surface of the pes. This is consistent also with younger averostran body fossils, which confirm an arthral arrangement of the digital pads. Among averostrans, squamous skin is confirmed in Ceratosauria (Carnotaurus), Allosauroidea (Allosaurus, Concavenator, Lourinhanosaurus), Compsognathidae (Juravenator), and Tyrannosauroidea (Santanaraptor, Albertosaurus, Daspletosaurus, Gorgosaurus, Tarbosaurus, Tyrannosaurus), whereas dermal ossifications consisting of sagittate and mosaic osteoderms are restricted to Ceratosaurus. Naked, non-scale bearing skin is found in the contentious tetanuran Sciurumimus, ornithomimosaurians (Ornithomimus) and possibly tyrannosauroids (Santanaraptor), and also on the patagia of scansoriopterygids (Ambopteryx, Yi). Scales are surprisingly conservative among non-avialan theropods compared to some dinosaurian groups (e.g. hadrosaurids); however, the limited preservation of tegument on most specimens hinders further interrogation. Scale patterns vary among and/or within body regions in Carnotaurus, Concavenator and Juravenator, and include polarised, snake-like ventral scales on the tail of the latter two genera. Unusual but more uniformly distributed patterning also occurs in Tyrannosaurus, whereas feature scales are present only in Albertosaurus and Carnotaurus. Few theropods currently show compelling evidence for the co-occurrence of scales and feathers (e.g. Juravenator, Sinornithosaurus), although reticulate scales were probably retained on the mani and pedes of many theropods with a heavy plumage. Feathers and filamentous structures appear to have replaced widespread scaly integuments in maniraptorans. Theropod skin, and that of dinosaurs more broadly, remains a virtually untapped area of study and the appropriation of commonly used techniques in other palaeontological fields to the study of skin holds great promise for future insights into the biology, taphonomy and relationships of these extinct animals.Fil: Hendrickx, Christophe Marie Fabian. Fundación Miguel Lillo; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico - Tucumán. Unidad Ejecutora Lillo; ArgentinaFil: Bell, Phil R.. University Of New England; AustraliaFil: Pittman, Michael. Colegio Universitario de Londres; Reino UnidoFil: Milner, Andrew R. C.. No especifíca;Fil: Cuesta Fidalgo, Elena. Bayerische Staatssammlung für Paläontologie und Geologie; AlemaniaFil: O'Connor, Jingmai. No especifíca;Fil: Loewen, Mark. No especifíca;Fil: Currie, Philip J.. University of Alberta; CanadáFil: Mateus, Octávio. Universidade Nova de Lisboa; PortugalFil: Kaye, Thomas G.. Foundation For Scientific Advancement; Estados UnidosFil: Delcourt, Rafael. Universidade Estadual de Campinas; Brasi

The scaly skin of the abelisaurid Carnotaurus sastrei (Theropoda: Ceratosauria) from the Upper Cretaceous of Patagonia

The integument of the theropod dinosaur Carnotaurus sastrei from the Upper Cretaceous of Argentina is here described in detail for the first time. The scaly skin of this abelisaurid is the most completely preserved of any theropod and the only example of this form of integument known outside of Tetanurae (excluding footprints). The skin is preserved in the shoulder, thoracic, tail and, possibly, neck regions and consists of medium to large (20–65 mm in diameter) conical feature scales surrounded by a network of low and small (1000 kg), we speculate that the skin may have played a vital role in thermoregulation; a role consistent with integument function in extant mammals and reptiles.Fil: Hendrickx, Christophe Marie Fabian. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico - Tucumán. Unidad Ejecutora Lillo; ArgentinaFil: Bell, Phil R.. University Of New England Australia; Australi

Crocodile-like sensory scales in a Late Jurassic theropod dinosaur

Early in amniote evolution, epidermal scales evolved in stem reptiles as an efficient barrier against water loss and ultraviolet radiation, making them a key development in the transition to a fully terrestrial existence [1]. Accordingly, epidermal scales are not simple inert structures but highly-evolved organs suited to perform a broad suite of functions. Here, we provide new data on the epidermal complexity of a non-avian theropod, Juravenator starki, from the Torleite Formation (upper Kimmeridgian), Bavaria, Germany [2]. Although epidermal scales have been noted previously on the tail of Juravenator, we report a unique scale type with distinctive circular nodes that we identify as integumentary sense organs, analogous to those in modern crocodylians. The surprising presence of such structures suggests the tail had a sensory function, which is nevertheless congruent with the inferred ecology of Juravenator and the evolution of integumentary sense organs among archosaurs. Bell and Hendrickx describe evidence for a crocodile-like sensory system of integumentary sense organs (ISOs) from fossilised soft tissues on the diminutive theropod dinosaur Juravenator from the Jurassic of Bavaria.Fil: Bell, Phil R.. University Of New England Australia; AustraliaFil: Hendrickx, Christophe Marie Fabian. Fundación Miguel Lillo; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Tucumán; Argentin