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      The African cynodont Aleodon (Cynodontia, Probainognathia) in the Triassic of southern Brazil and its biostratigraphic significance

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          Abstract

          In this contribution we report the first occurrence of the enigmatic African probainognathian genus Aleodon in the Middle-early Late Triassic of several localities from the state of Rio Grande do Sul in southern Brazil. Aleodon is unusual among early probainognathians in having transversely-expanded postcanine teeth, similar to those of gomphodont cynognathians. This genus was previously known from the Manda Beds of Tanzania and the upper Omingonde Formation of Namibia. The Brazilian record of this genus is based upon multiple specimens representing different ontogenetic stages, including three that were previously referred to the sectorial-toothed probainognathian Chiniquodon theotonicus. We propose a new species of Aleodon ( A. cromptoni sp. nov.) based on the specimens from Brazil. Additionally, we tentatively refer one specimen from the upper Omingonde Formation of Namibia to this new taxon, strengthening biostratigraphic correlations between these strata. Inclusion of A. cromptoni in a phylogenetic analysis of eucynodonts recovers it as the sister-taxon of A. brachyrhamphus within the family Chiniquodontidae. The discovery of numerous specimens of Aleodon among the supposedly monospecific Chiniquodon samples of Brazil raises concerns about chiniquodontid alpha taxonomy, particularly given the extremely broad geographic distribution of Chiniquodon. The discovery of Brazilian Aleodon and new records of the traversodontid Luangwa supports the hypothesis that at least two subzones can be recognized in the Dinodontosaurus Assemblage Zone.

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          Sequence stratigraphy of continental Triassic strata of Southernmost Brazil: a contribution to Southwestern Gondwana palaeogeography and palaeoclimate

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            Simplification as a trend in synapsid cranial evolution.

            The prevalence and meaning of morphological trends in the fossil record have undergone renewed scrutiny in recent years. Studies have typically focused on trends in body size evolution, which have yielded conflicting results, and have only rarely addressed the question as to whether other morphological characteristics show persistent directionality over long time scales. I investigated reduction in number of skull and lower jaw bones (through loss or fusion) over approximately 150 million years of premammalian synapsid history. The results of a new skull simplification metric (SSM), which is defined as a function of the number of distinct elements, show that pronounced simplification is evident on both temporal (i.e., stratigraphic) and phylogenetic scales. Postcranial evolution exhibits a similar pattern. Skull size, in contrast, bears little relationship with the number of distinct skull bones present. Synapsid skulls carried close to their observed maximum number of elements for most of the Late Carboniferous and Early Permian. The SSM decreased in the Late Permian but, coincident with the radiation of early therapsids, the range of observed SSM values widened during this interval. From derived nonmammalian cynodonts in the Early Triassic through the earliest mammals in the Early Jurassic, both the minimum and maximum SSM decreased. Data from three representative modern mammals (platypus, opossum, and human) suggest that this trend continues through the Cenozoic. In a phylogenetic context, the number of skull elements present in a taxon shows a significant negative relationship with the number of branching events passed from the root of the tree; more deeply embedded taxa have smaller SSM scores. This relationship holds for various synapsid subgroups as well. Although commonly ascribed to the effects of long-term selection, evolutionary trends can alternatively reflect an underlying intrinsic bias in morphological change. In the case of synapsid skull bones (and those of some other tetrapods lineages), the rare production of novel, or neomorphic, elements may have contributed to the observed trend toward skeletal simplification.
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              The radiation of cynodonts and the ground plan of mammalian morphological diversity

              Cynodont therapsids diversified extensively after the Permo-Triassic mass extinction event, and gave rise to mammals in the Jurassic. We use an enlarged and revised dataset of discrete skeletal characters to build a new phylogeny for all main cynodont clades from the Late Permian to the Early Jurassic, and we analyse models of morphological diversification in the group. Basal taxa and epicynodonts are paraphyletic relative to eucynodonts, and the latter are divided into cynognathians and probainognathians, with tritylodonts and mammals forming sister groups. Disparity analyses reveal a heterogeneous distribution of cynodonts in a morphospace derived from cladistic characters. Pairwise morphological distances are weakly correlated with phylogenetic distances. Comparisons of disparity by groups and through time are non-significant, especially after the data are rarefied. A disparity peak occurs in the Early/Middle Triassic, after which period the mean disparity fluctuates little. Cynognathians were characterized by high evolutionary rates and high diversity early in their history, whereas probainognathian rates were low. Community structure may have been instrumental in imposing different rates on the two clades.
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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
                14 June 2017
                2017
                : 12
                : 6
                : e0177948
                Affiliations
                [1 ]Laboratório de Paleontologia de Vertebrados, Departamento de Paleontologia e Estratigrafia, Instituto de Geociências, Universidade Federal do Rio Grande do Sul (UFRGS), Agronomia, Porto Alegre, Rio Grande do Sul, Brazil
                [2 ]Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Berlin, Germany
                [3 ]Seção de Paleontologia, Museu de Ciências Naturais da Fundação Zoobotânica do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, Brazil
                [4 ]Laboratório de Estratigrafia e Paleobiologia, Departamento de Geociências, Universidade Federal de Santa Maria, Santa Maria, Rio Grande do Sul, Brazil
                University of Michigan, UNITED STATES
                Author notes

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

                • Conceptualization: AGM.

                • Funding acquisition: AGM.

                • Investigation: AGM CFK TPM VDPN AMR AASD-R CLS MBS.

                • Writing – review & editing: AGM CFK TPM VDPN AMR AASD-R CLS MBS.

                Author information
                http://orcid.org/0000-0003-4489-0888
                Article
                PONE-D-16-48641
                10.1371/journal.pone.0177948
                5470689
                28614355
                cc0dff16-cbb9-4871-a57f-c98a66eba548
                © 2017 Martinelli et al

                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
                : 8 December 2016
                : 2 May 2017
                Page count
                Figures: 23, Tables: 3, Pages: 54
                Funding
                This work is supported by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and the Paleontological Society International Research Program (PalSIRP) - Sepkoski Grant (2016) to AGM. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
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