Untangling the dinosaur family tree

For over a century, the standard classification scheme has split dinosaurs into two fundamental groups: ‘lizard-hipped’ saurischians (including meat-eating theropods and long-necked sauropodomorphs) and ‘bird-hipped’ ornithischians (including a variety of herbivorous species).In a recent paper, Baron et al. challenged this paradigm with a new phylogenetic analysis that places theropods and ornithischians together in a group called Ornithoscelida, to the exclusion of sauropodomorphs, and used their phylogeny to argue that dinosaurs may have originated in northern Pangaea, not in the southern part of the supercontinent, as has more commonly been considered. Here we evaluate and reanalyse the morphological dataset underpinning the proposal by Baron et al. and provide quantitative biogeographic analyses, which challenge the key results of their study by recovering a classical monophyletic Saurischia and a Gondwanan origin for dinosaurs. This shows that the Ornithoscelida hypothesis is not the final word, and that there is still great uncertainty around the basic structure of the dinosaur family tree.Fil: Langer, Max C.. Universidade de Sao Paulo; BrasilFil: Ezcurra, Martin Daniel. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; ArgentinaFil: Rauhut, Oliver Walter Mischa. Ludwig Maximilians Universitat; AlemaniaFil: Benton, Michael J.. University of Bristol; Reino UnidoFil: Knoll, Fabien. University of Manchester; Reino UnidoFil: McPhee, Blair W.. Universidade de Sao Paulo; BrasilFil: Novas, Fernando Emilio. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; ArgentinaFil: Pol, Diego. Museo Paleontológico Egidio Feruglio; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; ArgentinaFil: Brusatte, Stephen L.. University of Edinburgh; Reino Unid

A new prozostrodontian cynodont (Therapsida) from the Late Triassic Riograndia Assemblage Zone (Santa Maria Supersequence) of Southern Brazil

We report here on a new prozostrodontian cynodont, Botucaraitherium belarminoi gen. et sp. nov., from the Late Triassic Riograndia Assemblage Zone (AZ) of the Candelária Sequence (Santa Maria Supersequence), collected in the Botucaraí Hill Site, Candelária Municipality, state of Rio Grande do Sul, Brazil. The new taxon is based on a single specimen (holotype MMACR-PV-003-T) which includes the left lower jaw, without postdentary bones, bearing the root of the last incisor, canine and four postcanines plus one partial crown inside the dentary, not erupted, and two maxillary fragments, one with a broken canine and another with one postcanine. The features of the lower jaw and lower/upper postcanines resemble those of the prozostrodontians Prozostrodon brasiliensis from the older Hyperodapedon AZ and Brasilodon quadrangularis and Brasilitherium riograndensis from the same Riograndia AZ. The inclusion of Botucaraitherium within a broad phylogenetic analysis, positioned it as a more derived taxon than tritylodontids, being the sister-taxon of Brasilodon, Brasilitherium plus Mammaliaformes. Although the new taxon is based on few cranial elements, it represents a additional faunal component of the Triassic Riograndia AZ of southern Brazil, in which small-sized derived non-mammaliaform cynodonts, closely related to the origin of mammaliaforms, were ecologically well succeed and taxonomically diverseNós reportamos aqui um novo cinodonte prozostrodonte, Botucaraitherium belarminoi gen. et sp. nov., do Triássico Tardio da Zona de Assembleia (ZA) de Riograndia da Sequência Candelária (Supersequência Santa Maria), coletado no afl oramento Sítio Botucaraí, no município de Candelária, Rio Grande do Sul, Brasil. O novo táxon está baseado em um único espécime (holótipo MMACR-PV- 003-T) o qual inclui a mandíbula esquerda, sem os ossos pós-dentários, com a raiz do último incisivo preservada, o canino e quatro dentes pós-caninos, além de uma coroa parcial, não erupcionada, do quinto pós-canino, e dois fragmentos maxilares, um com um canino quebrado, e outro portando apenas um dente pós-canino. As feições mandibulares e dentárias assemelham-se àquelas dos cinodontes prozostrodontes Prozostrodon brasiliensis da ZA de Hyperodapedon, mais antiga, e de Brasilodon quadrangularis e Brasilitherium riograndensis da mesma ZA de Riograndia. A inclusão de Botucaraitherium em uma ampla análise filogenética posicionou-o como um táxon mais derivado do que os tritilodontídeos, sendo o táxon-irmão de Brasilodon, Brasilitherium e mais Mammaliaformes. Apesar de o novo táxon ser baseado em poucos elementos cranianos, ele representa um componente faunístico adicional na ZA de Riograndia do Triássico sul-brasileiro, na qual os cinodontes não-mamaliaformes de pequeno tamanho, intimamente relacionados à origem dos mamíferos, foram ecologicamente bem sucedidos e taxonomicamente diverso

Deep evolutionary diversification of semicircular canals in archosaurs

Living archosaurs (birds and crocodylians) have disparate locomotor strategies that evolved since their divergence ∼250 mya. Little is known about the early evolution of the sensory structures that are coupled with these changes, mostly due to limited sampling of early fossils on key stem lineages. In particular, the morphology of the semicircular canals (SCCs) of the endosseous labyrinth has a long-hypothesized relationship with locomotion. Here, we analyze SCC shapes and sizes of living and extinct archosaurs encompassing diverse locomotor habits, including bipedal, semi-aquatic, and flying taxa. We test form-function hypotheses of the SCCs and chronicle their evolution during deep archosaurian divergences. We find that SCC shape is statistically associated with both flight and bipedalism. However, this shape variation is small and is more likely explained by changes in braincase geometry than by locomotor changes. We demonstrate high disparity of both shape and size among stem-archosaurs and a deep divergence of SCC morphologies at the bird–crocodylian split. Stem-crocodylians exhibit diverse morphologies, including aspects also present in birds and distinct from other reptiles. Therefore, extant crocodylian SCC morphologies do not reflect retention of a “primitive” reptilian condition. Key aspects of bird SCC morphology that hitherto were interpreted as flight related, including large SCC size and enhanced sensitivity, appeared early on the bird stem-lineage in non-flying dinosaur precursors. Taken together, our results indicate a deep divergence of SCC traits at the bird–crocodylian split and that living archosaurs evolved from an early radiation with high sensory diversity

Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria

Pterosaurs were the first vertebrates to evolve powered flight1 and comprised one of the main evolutionary radiations in terrestrial ecosystems of the Mesozoic era (approximately 252–66 million years ago), but their origin has remained an unresolved enigma in palaeontology since the nineteenth century2,3,4. These flying reptiles have been hypothesized to be the close relatives of a wide variety of reptilian clades, including dinosaur relatives2,3,4,5,6,7,8, and there is still a major morphological gap between those forms and the oldest, unambiguous pterosaurs from the Upper Triassic series. Here, using recent discoveries of well-preserved cranial remains, microcomputed tomography scans of fragile skull bones (jaws, skull roofs and braincases) and reliably associated postcrania, we demonstrate that lagerpetids—a group of cursorial, non-volant dinosaur precursors—are the sister group of pterosaurs, sharing numerous synapomorphies across the entire skeleton. This finding substantially shortens the temporal and morphological gap between the oldest pterosaurs and their closest relatives and simultaneously strengthens the evidence that pterosaurs belong to the avian line of archosaurs. Neuroanatomical features related to the enhanced sensory abilities of pterosaurs9 are already present in lagerpetids, which indicates that these features evolved before flight. Our evidence illuminates the first steps of the assembly of the pterosaur body plan, whose conquest of aerial space represents a remarkable morphofunctional innovation in vertebrate evolution

Periodontal Ligament, Cementum, and Alveolar Bone in the Oldest Herbivorous Tetrapods, and Their Evolutionary Significance

Tooth implantation provides important phylogenetic and functional information about the dentitions of amniotes. Traditionally, only mammals and crocodilians have been considered truly thecodont, because their tooth roots are coated in layers of cementum for anchorage of the periodontal ligament, which is in turn attached to the bone lining the alveolus, the alveolar bone. The histological properties and developmental origins of these three periodontal tissues have been studied extensively in mammals and crocodilians, but the identities of the periodontal tissues in other amniotes remain poorly studied. Early work on dental histology of basal amniotes concluded that most possess a simplified tooth attachment in which the tooth root is ankylosed to a pedestal composed of “bone of attachment”, which is in turn fused to the jaw. More recent studies have concluded that stereotypically thecodont tissues are also present in non-mammalian, non-crocodilian amniotes, but these studies were limited to crown groups or secondarily aquatic reptiles. As the sister group to Amniota, and the first tetrapods to exhibit dental occlusion, diadectids are the ideal candidates for studies of dental evolution among terrestrial vertebrates because they can be used to test hypotheses of development and homology in deep time. Our study of Permo-Carboniferous diadectid tetrapod teeth and dental tissues reveal the presence of two types of cementum, periodontal ligament, and alveolar bone, and therefore the earliest record of true thecodonty in a tetrapod. These discoveries in a stem amniote allow us to hypothesize that the ability to produce the tissues that characterize thecodonty in mammals and crocodilians is very ancient and plesiomorphic for Amniota. Consequently, all other forms of tooth implantation in crown amniotes are derived arrangements of one or more of these periodontal tissues and not simply ankylosis of teeth to the jaw by plesiomorphically retaining “bone of attachment”, as previously suggested