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      Synchrotron Reveals Early Triassic Odd Couple: Injured Amphibian and Aestivating Therapsid Share Burrow

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          Abstract

          Fossorialism is a beneficial adaptation for brooding, predator avoidance and protection from extreme climate. The abundance of fossilised burrow casts from the Early Triassic of southern Africa is viewed as a behavioural response by many tetrapods to the harsh conditions following the Permo-Triassic mass-extinction event. However, scarcity of vertebrate remains associated with these burrows leaves many ecological questions unanswered. Synchrotron scanning of a lithified burrow cast from the Early Triassic of the Karoo unveiled a unique mixed-species association: an injured temnospondyl amphibian ( Broomistega) that sheltered in a burrow occupied by an aestivating therapsid ( Thrinaxodon). The discovery of this rare rhinesuchid represents the first occurrence in the fossil record of a temnospondyl in a burrow. The amphibian skeleton shows signs of a crushing trauma with partially healed fractures on several consecutive ribs. The presence of a relatively large intruder in what is interpreted to be a Thrinaxodon burrow implies that the therapsid tolerated the amphibian’s presence. Among possible explanations for such unlikely cohabitation, Thrinaxodon aestivation is most plausible, an interpretation supported by the numerous Thrinaxodon specimens fossilised in curled-up postures. Recent advances in synchrotron imaging have enabled visualization of the contents of burrow casts, thus providing a novel tool to elucidate not only anatomy but also ecology and biology of ancient tetrapods.

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          A review of burrowing by semi-fossorial vertebrates in arid environments

          Al Kinlaw (1999)
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            The endocast of MH1, Australopithecus sediba.

            The virtual endocast of MH1 (Australopithecus sediba), obtained from high-quality synchrotron scanning, reveals generally australopith-like convolutional patterns on the frontal lobes but also some foreshadowing of features of the human frontal lobes, such as posterior repositioning of the olfactory bulbs. Principal component analysis of orbitofrontal dimensions on australopith endocasts (MH1, Sts 5, and Sts 60) indicates that among these, MH1 orbitofrontal shape and organization align most closely with human endocasts. These results are consistent with gradual neural reorganization of the orbitofrontal region in the transition from Australopithecus to Homo, but given the small volume of the MH1 endocast, they are not consistent with gradual brain enlargement before the transition.
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              Hibernation and daily torpor minimize mammalian extinctions.

              Small mammals appear to be less vulnerable to extinction than large species, but the underlying reasons are poorly understood. Here, we provide evidence that almost all (93.5%) of 61 recently extinct mammal species were homeothermic, maintaining a constant high body temperature and thus energy expenditure, which demands a high intake of food, long foraging times, and thus exposure to predators. In contrast, only 6.5% of extinct mammals were likely heterothermic and employed multi-day torpor (hibernation) or daily torpor, even though torpor is widespread within more than half of all mammalian orders. Torpor is characterized by substantial reductions of body temperature and energy expenditure and enhances survival during adverse conditions by minimizing food and water requirements, and consequently reduces foraging requirements and exposure to predators. Moreover, because life span is generally longer in heterothermic mammals than in related homeotherms, heterotherms can employ a 'sit-and-wait' strategy to withstand adverse periods and then repopulate when circumstances improve. Thus, torpor is a crucial but hitherto unappreciated attribute of small mammals for avoiding extinction. Many opportunistic heterothermic species, because of their plastic energetic requirements, may also stand a better chance of future survival than homeothermic species in the face of greater climatic extremes and changes in environmental conditions caused by global warming.
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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
                2013
                21 June 2013
                : 8
                : 6
                : e64978
                Affiliations
                [1 ]Evolutionary Studies Institute, University of the Witwatersrand, Johannesburg, Gauteng, South Africa
                [2 ]Department of Anthropology, Indiana University, Bloomington, Indiana, United States of America
                [3 ]Museum of Central Australia, Araluen Cultural Precinct, Alice Springs, Northern Territory, Australia
                [4 ]European Synchrotron Radiation Facility, Grenoble, France
                Ludwig-Maximilians-Universität München, Germany
                Author notes

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

                Conceived and designed the experiments: VF KJC BSR PT. Performed the experiments: VF PT. Analyzed the data: VF FA KJC DCC BSR AY PT. Contributed reagents/materials/analysis tools: VF PT. Wrote the paper: VF FA KJC DCC BSR PT.

                Article
                PONE-D-12-38038
                10.1371/journal.pone.0064978
                3689844
                23805181
                80fdc32c-d21e-4d61-a167-26476f19ddbc
                Copyright @ 2013

                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
                : 4 December 2012
                : 18 April 2013
                Page count
                Pages: 7
                Funding
                This work was supported by the ESRF (proposal EC-847), the University Research Committee (URC) of Wits, and the Palaeontological Scientific Trust (PAST). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology
                Ecology
                Behavioral Ecology
                Community Ecology
                Paleoecology
                Evolutionary Biology
                Behavioral Ecology
                Earth Sciences
                Geology
                Stratigraphy
                Lithostratigraphy
                Sedimentary Geology
                Paleontology
                Paleobiology
                Paleoecology
                Taphonomy
                Vertebrate Paleontology

                Uncategorized
                Uncategorized

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