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      Cancellous bone and theropod dinosaur locomotion. Part I—an examination of cancellous bone architecture in the hindlimb bones of theropods

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          Abstract

          This paper is the first of a three-part series that investigates the architecture of cancellous (‘spongy’) bone in the main hindlimb bones of theropod dinosaurs, and uses cancellous bone architectural patterns to infer locomotor biomechanics in extinct non-avian species. Cancellous bone is widely known to be highly sensitive to its mechanical environment, and has previously been used to infer locomotor biomechanics in extinct tetrapod vertebrates, especially primates. Despite great promise, cancellous bone architecture has remained little utilized for investigating locomotion in many other extinct vertebrate groups, such as dinosaurs. Documentation and quantification of architectural patterns across a whole bone, and across multiple bones, can provide much information on cancellous bone architectural patterns and variation across species. Additionally, this also lends itself to analysis of the musculoskeletal biomechanical factors involved in a direct, mechanistic fashion.

          On this premise, computed tomographic and image analysis techniques were used to describe and analyse the three-dimensional architecture of cancellous bone in the main hindlimb bones of theropod dinosaurs for the first time. A comprehensive survey across many extant and extinct species is produced, identifying several patterns of similarity and contrast between groups. For instance, more stemward non-avian theropods (e.g. ceratosaurs and tyrannosaurids) exhibit cancellous bone architectures more comparable to that present in humans, whereas species more closely related to birds (e.g. paravians) exhibit architectural patterns bearing greater similarity to those of extant birds. Many of the observed patterns may be linked to particular aspects of locomotor biomechanics, such as the degree of hip or knee flexion during stance and gait. A further important observation is the abundance of markedly oblique trabeculae in the diaphyses of the femur and tibia of birds, which in large species produces spiralling patterns along the endosteal surface. Not only do these observations provide new insight into theropod anatomy and behaviour, they also provide the foundation for mechanistic testing of locomotor hypotheses via musculoskeletal biomechanical modelling.

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          A Guideline of Selecting and Reporting Intraclass Correlation Coefficients for Reliability Research.

          Intraclass correlation coefficient (ICC) is a widely used reliability index in test-retest, intrarater, and interrater reliability analyses. This article introduces the basic concept of ICC in the content of reliability analysis.
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            Intraclass correlations: Uses in assessing rater reliability.

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              The global diversity of birds in space and time.

              Current global patterns of biodiversity result from processes that operate over both space and time and thus require an integrated macroecological and macroevolutionary perspective. Molecular time trees have advanced our understanding of the tempo and mode of diversification and have identified remarkable adaptive radiations across the tree of life. However, incomplete joint phylogenetic and geographic sampling has limited broad-scale inference. Thus, the relative prevalence of rapid radiations and the importance of their geographic settings in shaping global biodiversity patterns remain unclear. Here we present, analyse and map the first complete dated phylogeny of all 9,993 extant species of birds, a widely studied group showing many unique adaptations. We find that birds have undergone a strong increase in diversification rate from about 50 million years ago to the near present. This acceleration is due to a number of significant rate increases, both within songbirds and within other young and mostly temperate radiations including the waterfowl, gulls and woodpeckers. Importantly, species characterized with very high past diversification rates are interspersed throughout the avian tree and across geographic space. Geographically, the major differences in diversification rates are hemispheric rather than latitudinal, with bird assemblages in Asia, North America and southern South America containing a disproportionate number of species from recent rapid radiations. The contribution of rapidly radiating lineages to both temporal diversification dynamics and spatial distributions of species diversity illustrates the benefits of an inclusive geographical and taxonomical perspective. Overall, whereas constituent clades may exhibit slowdowns, the adaptive zone into which modern birds have diversified since the Cretaceous may still offer opportunities for diversification.
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                Author and article information

                Contributors
                Journal
                PeerJ
                PeerJ
                PeerJ
                PeerJ
                PeerJ
                PeerJ Inc. (San Diego, USA )
                2167-8359
                31 October 2018
                2018
                : 6
                : e5778
                Affiliations
                [1 ]Geosciences Program, Queensland Museum , Brisbane, QLD, Australia
                [2 ]School of Allied Health Sciences, Griffith University , Gold Coast, QLD, Australia
                [3 ]Gold Coast Orthopaedic Research, Engineering and Education Alliance, Menzies Health Institute Queensland , Gold Coast, QLD, Australia
                [4 ]Current affiliation: Structure and Motion Laboratory, Department of Comparative Biomedical Sciences, Royal Veterinary College , Hatfield, Hertfordshire, UK
                [5 ]School of Biosciences, University of Melbourne , Melbourne, VIC, Australia
                [6 ]School of Science and Engineering, University of the Sunshine Coast , Maroochydore, QLD, Australia
                [7 ]School of Biological Sciences, University of Queensland , Brisbane, QLD, Australia
                [8 ]Structure and Motion Laboratory, Department of Comparative Biomedical Sciences, Royal Veterinary College , Hatfield, Hertfordshire, UK
                [9 ]Raymond M. Alf Museum of Paleontology at The Webb Schools , Claremont, CA, USA
                [10 ]Exercise and Human Performance, Menzies Health Institute Queensland , Gold Coast, QLD, Australia
                Author information
                http://orcid.org/0000-0003-4157-8434
                http://orcid.org/0000-0001-8174-3890
                http://orcid.org/0000-0002-6767-7038
                http://orcid.org/0000-0002-6930-2002
                http://orcid.org/0000-0003-1295-6395
                http://orcid.org/0000-0002-0824-9682
                Article
                5778
                10.7717/peerj.5778
                6215452
                30402347
                32332f3f-e95c-42fb-85bb-feb12ab69b76
                © 2018 Bishop et al.

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, reproduction and adaptation in any medium and for any purpose provided that it is properly attributed. For attribution, the original author(s), title, publication source (PeerJ) and either DOI or URL of the article must be cited.

                History
                : 8 January 2018
                : 18 September 2018
                Funding
                Funded by: An Australian Government Research Training Program Scholarship
                Funded by: The Paleontological Society
                Funded by: An International Society of Biomechanics Matching Dissertation Grant
                Funded by: An Australian Research Council DECRA Fellowship
                Award ID: DE120101503
                Funded by: The donation of CT scan time and technical assistance by Queensland X-ray
                This study was financially supported by an Australian Government Research Training Program Scholarship (to Peter Bishop), the Paleontological Society (Robert J. Stanton & James R. Dodd Award, to Peter Bishop), an International Society of Biomechanics Matching Dissertation Grant (to Peter Bishop), an Australian Research Council DECRA Fellowship (DE120101503, to Christofer Clemente) and the donation of CT scan time and technical assistance by Queensland X-ray (to Scott Hocknull). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Evolutionary Studies
                Paleontology
                Zoology

                cancellous bone,theropod,bird,locomotion,biomechanics
                cancellous bone, theropod, bird, locomotion, biomechanics

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