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      The Cranial Osteology and Feeding Ecology of the Metriorhynchid Crocodylomorph Genera Dakosaurus and Plesiosuchus from the Late Jurassic of Europe

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          Abstract

          Background

          Dakosaurus and Plesiosuchus are characteristic genera of aquatic, large-bodied, macrophagous metriorhynchid crocodylomorphs. Recent studies show that these genera were apex predators in marine ecosystems during the latter part of the Late Jurassic, with robust skulls and strong bite forces optimized for feeding on large prey.

          Methodology/Principal Findings

          Here we present comprehensive osteological descriptions and systematic revisions of the type species of both genera, and in doing so we resurrect the genus Plesiosuchus for the species Dakosaurus manselii. Both species are diagnosed with numerous autapomorphies. Dakosaurus maximus has premaxillary ‘lateral plates’; strongly ornamented maxillae; macroziphodont dentition; tightly fitting tooth-to-tooth occlusion; and extensive macrowear on the mesial and distal margins. Plesiosuchus manselii is distinct in having: non-amblygnathous rostrum; long mandibular symphysis; microziphodont teeth; tooth-crown apices that lack spalled surfaces or breaks; and no evidence for occlusal wear facets. Our phylogenetic analysis finds Dakosaurus maximus to be the sister taxon of the South American Dakosaurus andiniensis, and Plesiosuchus manselii in a polytomy at the base of Geosaurini (the subclade of macrophagous metriorhynchids that includes Dakosaurus, Geosaurus and Torvoneustes).

          Conclusions/Significance

          The sympatry of Dakosaurus and Plesiosuchus is curiously similar to North Atlantic killer whales, which have one larger ‘type’ that lacks tooth-crown breakage being sympatric with a smaller ‘type’ that has extensive crown breakage. Assuming this morphofunctional complex is indicative of diet, then Plesiosuchus would be a specialist feeding on other marine reptiles while Dakosaurus would be a generalist and possible suction-feeder. This hypothesis is supported by Plesiosuchus manselii having a very large optimum gape (gape at which multiple teeth come into contact with a prey-item), while Dakosaurus maximus possesses craniomandibular characteristics observed in extant suction-feeding odontocetes: shortened tooth-row, amblygnathous rostrum and a very short mandibular symphysis. We hypothesise that trophic specialisation enabled these two large-bodied species to coexist in the same ecosystem.

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          Most cited references7

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          Ecological, morphological and genetic divergence of sympatric North Atlantic killer whale populations.

          Ecological divergence has a central role in speciation and is therefore an important source of biodiversity. Studying the micro-evolutionary processes of ecological diversification at its early stages provides an opportunity for investigating the causative mechanisms and ecological conditions promoting divergence. Here we use morphological traits, nitrogen stable isotope ratios and tooth wear to characterize two disparate types of North Atlantic killer whale. We find a highly specialist type, which reaches up to 8.5 m in length and a generalist type which reaches up to 6.6 m in length. There is a single fixed genetic difference in the mtDNA control region between these types, indicating integrity of groupings and a shallow divergence. Phylogenetic analysis indicates this divergence is independent of similar ecological divergences in the Pacific and Antarctic. Niche-width in the generalist type is more strongly influenced by between-individual variation rather than within-individual variation in the composition of the diet. This first step to divergent specialization on different ecological resources provides a rare example of the ecological conditions at the early stages of adaptive radiation.
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            Shape and mechanics in thalattosuchian (Crocodylomorpha) skulls: implications for feeding behaviour and niche partitioning.

            Variation in modern crocodilian and extinct thalattosuchian crocodylomorph skull morphology is only weakly correlated with phylogeny, implying that factors other than evolutionary proximity play important roles in determining crocodile skull shape. To further explore factors potentially influencing morphological differentiation within the Thalattosuchia, we examine teleosaurid and metriorhynchid skull shape variation within a mechanical and dietary context using a combination of finite element modelling and multivariate statistics. Patterns of stress distribution through the skull were found to be very similar in teleosaurid and metriorhynchid species, with stress peaking at the posterior constriction of the snout and around the enlarged supratemporal fenestrae. However, the magnitudes of stresses differ, with metriorhynchids having generally stronger skulls. As with modern crocodilians, a strong linear relationship between skull length and skull strength exists, with short-snouted morphotypes experiencing less stress through the skull than long-snouted morphotypes under equivalent loads. Selection on snout shape related to dietary preference was found to work in orthogonal directions in the two families: diet is associated with snout length in teleosaurids and with snout width in metriorhynchids, suggesting that teleosaurid skulls were adapted for speed of attack and metriorhynchid skulls for force production. Evidence also indicates that morphological and functional differentiation of the skull occurred as a result of dietary preference, allowing closely related sympatric species to exploit a limited environment. Comparisons of the mechanical performance of the thalattosuchian skull with extant crocodilians show that teleosaurids and long-snouted metriorhynchids exhibit stress magnitudes similar to or greater than those of long-snouted modern forms, whereas short-snouted metriorhynchids display stress magnitudes converging on those found in short-snouted modern species. As a result, teleosaurids and long-snouted metriorhynchids were probably restricted to lateral attacks of the head and neck, but short-snouted metriorhynchids may have been able to employ the grasp and shake and/or 'death roll' feeding and foraging behaviours.
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              Salt glands in the Jurassic metriorhynchid Geosaurus: implications for the evolution of osmoregulation in Mesozoic marine crocodyliforms.

              The presence of salt-excreting glands in extinct marine sauropsids has been long suspected based on skull morphology. Previously, we described for the first time the natural casts of salt-excreting glands in the head of the Jurassic metriorhynchid crocodyliform Geosaurus araucanensis from the Tithonian of the Vaca Muerta Formation in the Neuquén Basin (Argentina). In the present study, salt-excreting glands are identified in three new individuals (adult, a sub-adult and a juvenile) referable to the same species. New material provides significant information on the salt glands form and function and permit integration of evolutionary scenarios proposed on a physiological basis in extant taxa with evidence from the fossil record. G. araucanensis represents an advanced stage of the basic physiological model to marine adaptations in reptiles. G. araucanensis salt glands were hypertrophied. On this basis, it can be hypothesized that these glands had a high excretory capability. This stage implies that G. araucanensis (like extant pelagic reptiles, e.g. cheloniids) could have maintained constant plasma osmolality even when seawater or osmoconforming prey were ingested. A gradual model of marine adaptation in crocodyliforms based on physiology (freshwater to coastal/estuarine to estuarine /marine to pelagic life) is congruent with the phylogeny of crocodyliforms based on skeletal morphology. The fossil record suggests that the stage of marine pelagic adaptation was achieved by the Early Middle Jurassic. Salt gland size in the juvenile suggests that juveniles were, like adults, pelagic.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2012
                18 September 2012
                : 7
                : 9
                : e44985
                Affiliations
                [1 ]School of Geosciences, University of Edinburgh, Edinburgh, United Kingdom
                [2 ]Division of Paleontology, American Museum of Natural History, New York City, New York, United States of America
                [3 ]Department of Earth and Environmental Sciences, Columbia University, New York City, New York, United States of America
                [4 ]Departamento de Paleontologia e Estratigrafia, Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil
                [5 ]División de Paleontología, Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’, Buenos Aires, Argentina
                [6 ]Department of Anatomy, New York College of Osteopathic Medicine, Old Westbury, New York, United States of America
                [7 ]Department of Earth Sciences, Natural History Museum, London, United Kingdom
                [8 ]Departamento Paleontología de Vertebrados, Museo de La Plata, La Plata, Argentina
                [9 ]School of Earth Sciences, University of Bristol, Bristol, United Kingdom
                [10 ]Museo del Jurásico de Asturias (MUJA), Colunga, Spain
                [11 ]Departamento de Geología, Universidad de Oviedo, Oviedo, Spain
                [12 ]Staatliches Museum für Naturkunde, Stuttgart, Germany
                Ludwig-Maximilians-Universität München, Germany
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: MTY. Performed the experiments: MTY MS. Analyzed the data: MTY SLB BLB MS JIR-O. Contributed reagents/materials/analysis tools: MBdA JBD LS MSF JIR-O RRS. Wrote the paper: MTY SLB BLB JIR-O.

                Article
                PONE-D-12-11584
                10.1371/journal.pone.0044985
                3445579
                23028723
                27ad216a-d5d2-40c8-a143-6840ae611512

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 24 April 2012
                : 16 August 2012
                Page count
                Pages: 42
                Funding
                The authors have no support or funding to report.
                Categories
                Research Article
                Biology
                Evolutionary Biology
                Evolutionary Systematics
                Taxonomy
                Animal Taxonomy
                Cladistics
                Paleontology
                Paleobiology
                Zoology
                Animal Taxonomy
                Comparative Anatomy
                Earth Sciences
                Paleontology
                Paleobiology
                Vertebrate Paleontology

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                Uncategorized

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