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      Craniodental and humeral morphology of a new species of Masrasector (Teratodontinae, Hyaenodonta, Placentalia) from the late Eocene of Egypt and locomotor diversity in hyaenodonts

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          Abstract

          Hyaenodonta is a diverse clade of carnivorous mammals that were part of terrestrial faunas in the Paleogene of Eurasia and North America, but the oldest record for the group is Afro-Arabian, making the record there vital for understanding the evolution of this wide-spread group. Previous studies show an ancient split between two major clades of hyaenodonts that converged in hypercarnivory: Hyainailourinae and Hyaenodontinae. These clades are each supported by cranial characters. Phylogenetic analyses of hyaenodonts also support the monophyly of Teratodontinae, an Afro-Arabian clade of mesocarnivorous to hypercarnivorous hyaenodonts. Unfortunately, the cranial anatomy of teratodontines is poorly known, and aligning the clade with other lineages has been difficult. Here, a new species of the phylogenetically controversial teratodontine Masrasector is described from Locality 41 (latest Priabonian, late Eocene) from the Fayum Depression, Egypt. The hypodigm includes the most complete remains of a Paleogene teratodontine, including largely complete crania, multiple dentaries, and isolated humeri. Standard and “tip-dating” Bayesian analyses of a character-taxon matrix that samples cranial, postcranial, and dental characters support a monophyletic Masrasector within Teratodontinae, which is consistently placed as a close sister group of Hyainailouridae. The cranial morphology of Masrasector provides new support for an expanded Hyainailouroidea (Teratodontinae + Hyainailouridae), particularly characters of the nuchal crest, palate, and basicranium. A discriminant function analysis was performed using measurements of the distal humerus from a diverse sample of extant carnivorans to infer the locomotor habits of Masrasector. Masrasector was assigned to the “terrestrial” locomotor category, a result consistent with the well-defined medial trochlear ridges, and moderately developed supinator crests of the specimens. Masrasector appears to have been a fast-moving terrestrial form with a diverse diet. These specimens considerably improve our understanding of Teratodontinae, an ancient member of the Afro-Arabian mammalian fauna, and our understanding of hyaenodont diversity before the dispersal of Carnivora to the continent near the end of the Paleogene.

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          NIH Image to ImageJ: 25 years of image analysis.

          For the past 25 years NIH Image and ImageJ software have been pioneers as open tools for the analysis of scientific images. We discuss the origins, challenges and solutions of these two programs, and how their history can serve to advise and inform other software projects.
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            To discover interordinal relationships of living and fossil placental mammals and the time of origin of placentals relative to the Cretaceous-Paleogene (K-Pg) boundary, we scored 4541 phenomic characters de novo for 86 fossil and living species. Combining these data with molecular sequences, we obtained a phylogenetic tree that, when calibrated with fossils, shows that crown clade Placentalia and placental orders originated after the K-Pg boundary. Many nodes discovered using molecular data are upheld, but phenomic signals overturn molecular signals to show Sundatheria (Dermoptera + Scandentia) as the sister taxon of Primates, a close link between Proboscidea (elephants) and Sirenia (sea cows), and the monophyly of echolocating Chiroptera (bats). Our tree suggests that Placentalia first split into Xenarthra and Epitheria; extinct New World species are the oldest members of Afrotheria.
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              A general comparison of relaxed molecular clock models.

              Several models have been proposed to relax the molecular clock in order to estimate divergence times. However, it is unclear which model has the best fit to real data and should therefore be used to perform molecular dating. In particular, we do not know whether rate autocorrelation should be considered or which prior on divergence times should be used. In this work, we propose a general bench mark of alternative relaxed clock models. We have reimplemented most of the already existing models, including the popular lognormal model, as well as various prior choices for divergence times (birth-death, Dirichlet, uniform), in a common Bayesian statistical framework. We also propose a new autocorrelated model, called the "CIR" process, with well-defined stationary properties. We assess the relative fitness of these models and priors, when applied to 3 different protein data sets from eukaryotes, vertebrates, and mammals, by computing Bayes factors using a numerical method called thermodynamic integration. We find that the 2 autocorrelated models, CIR and lognormal, have a similar fit and clearly outperform uncorrelated models on all 3 data sets. In contrast, the optimal choice for the divergence time prior is more dependent on the data investigated. Altogether, our results provide useful guidelines for model choice in the field of molecular dating while opening the way to more extensive model comparisons.
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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, CA USA )
                1932-6203
                19 April 2017
                2017
                : 12
                : 4
                : e0173527
                Affiliations
                [1 ]Department of Biomedical Sciences, Heritage College of Osteopathic Medicine, Ohio University, Athens, Ohio, United States of America
                [2 ]Department of Integrative Anatomical Sciences, Keck School of Medicine, University of Southern California, Los Angeles, California, United States of America
                Institute of Vertebrate Paleontology and Paleoanthropology Chinese Academy of Sciences, CHINA
                Author notes

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

                • Conceptualization: MRB ERS.

                • Data curation: MRB.

                • Formal analysis: MRB ERS.

                • Funding acquisition: MRB ERS.

                • Investigation: MRB ERS.

                • Methodology: MRB ERS.

                • Project administration: MRB.

                • Resources: MRB ERS.

                • Software: MRB ERS.

                • Supervision: ERS.

                • Validation: MRB ERS.

                • Visualization: MRB.

                • Writing – original draft: MRB.

                • Writing – review & editing: ERS.

                Author information
                http://orcid.org/0000-0002-3840-5308
                Article
                PONE-D-16-43912
                10.1371/journal.pone.0173527
                5396875
                28422967
                035b3f57-8905-4d43-9a52-ac6101357c10
                © 2017 Borths, Seiffert

                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 November 2016
                : 17 February 2017
                Page count
                Figures: 19, Tables: 8, Pages: 60
                Funding
                Funded by: funder-id http://dx.doi.org/10.13039/100000001, National Science Foundation;
                Award ID: BCS-0416164
                Award Recipient :
                Funded by: funder-id http://dx.doi.org/10.13039/100000169, Division of Behavioral and Cognitive Sciences;
                Award ID: BCS-0819186
                Award Recipient :
                Funded by: funder-id http://dx.doi.org/10.13039/100000169, Division of Behavioral and Cognitive Sciences;
                Award ID: BCS-1231288
                Award Recipient :
                Funded by: Leakey Foundation (US)
                Award Recipient :
                Funded by: Gordon and Ann Getty
                Award Recipient :
                Funded by: funder-id http://dx.doi.org/10.13039/100000155, Division of Environmental Biology;
                Award ID: DEB-1311354
                Award Recipient : Matthew Robert Borths
                Funded by: The Explorers Club
                Award Recipient : Matthew Robert Borths
                Funded by: funder-id http://dx.doi.org/10.13039/100000076, Directorate for Biological Sciences;
                Award ID: DBI 1612062
                Award Recipient : Matthew Robert Borths
                Funded by: Society for Integrative and Comparative Biology
                Award ID: Graduate Student Travel Award
                Award Recipient : Matthew Robert Borths
                Field work in the Fayum Depression, Egypt, and digital curation of Fayum fossils is supported by the U.S. National Science Foundation through grants BCS-0416164 ( https://www.nsf.gov/awardsearch/showAward?AWD_ID=0416164), BCS-0819186 ( https://nsf.gov/awardsearch/showAward?AWD_ID=0819186), and BCS-1231288 ( https://www.nsf.gov/awardsearch/showAward?AWD_ID=1231288), Gordon and Ann Getty, and The Leakey Foundation ( https://leakeyfoundation.org/). MRB was supported by a U.S. National Science Foundation Doctoral Dissertation Improvement Grant (DEB-1311354; https://www.nsf.gov/awardsearch/showAward?AWD_ID=1311354&HistoricalAwards=false), a National Science Foundation Graduate Research Fellowship ( https://www.nsfgrfp.org/), a Turkana Basin Institute Graduate Fellowship ( http://www.turkanabasin.org/about/fellowships/), and a Stony Brook University Graduate Council Fellowship, and is currently supported by a U.S. National Science Foundation Postdoctoral Fellowship in Biology (DBI-1612062; https://www.nsf.gov/awardsearch/showAward?AWD_ID=1612062). Portions of the data acquisition for this study were also supported by grants from The Explorers Club ( https://explorers.org/expeditions/funding/expedition_grants) and the Society for Integrative and Comparative Biology ( http://www.sicb.org/grants/fgstinfo.php). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
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                Custom metadata
                Data is available from MorphoSource with the identifier P191: Craniodental and humeral morphology of a new species of *Masrasector *(Teratodontinae, Hyaenodonta, Placentalia) from the late Eocene of Egypt and locomotor diversity in hyaenodonts.

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