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      Improved Phylogenomic Taxon Sampling Noticeably Affects Nonbilaterian Relationships

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          Abstract

          Despite expanding data sets and advances in phylogenomic methods, deep-level metazoan relationships remain highly controversial. Recent phylogenomic analyses depart from classical concepts in recovering ctenophores as the earliest branching metazoan taxon and propose a sister-group relationship between sponges and cnidarians (e.g., Dunn CW, Hejnol A, Matus DQ, et al. (18 co-authors). 2008. Broad phylogenomic sampling improves resolution of the animal tree of life. Nature 452:745–749). Here, we argue that these results are artifacts stemming from insufficient taxon sampling and long-branch attraction (LBA). By increasing taxon sampling from previously unsampled nonbilaterians and using an identical gene set to that reported by Dunn et al., we recover monophyletic Porifera as the sister group to all other Metazoa. This suggests that the basal position of the fast-evolving Ctenophora proposed by Dunn et al. was due to LBA and that broad taxon sampling is of fundamental importance to metazoan phylogenomic analyses. Additionally, saturation in the Dunn et al. character set is comparatively high, possibly contributing to the poor support for some nonbilaterian nodes.

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          The Trichoplax genome and the nature of placozoans.

          As arguably the simplest free-living animals, placozoans may represent a primitive metazoan form, yet their biology is poorly understood. Here we report the sequencing and analysis of the approximately 98 million base pair nuclear genome of the placozoan Trichoplax adhaerens. Whole-genome phylogenetic analysis suggests that placozoans belong to a 'eumetazoan' clade that includes cnidarians and bilaterians, with sponges as the earliest diverging animals. The compact genome shows conserved gene content, gene structure and synteny in relation to the human and other complex eumetazoan genomes. Despite the apparent cellular and organismal simplicity of Trichoplax, its genome encodes a rich array of transcription factor and signalling pathway genes that are typically associated with diverse cell types and developmental processes in eumetazoans, motivating further searches for cryptic cellular complexity and/or as yet unobserved life history stages.
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            Generelle Morphologie der Organismen

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              Phylogenomics revives traditional views on deep animal relationships.

              The origin of many of the defining features of animal body plans, such as symmetry, nervous system, and the mesoderm, remains shrouded in mystery because of major uncertainty regarding the emergence order of the early branching taxa: the sponge groups, ctenophores, placozoans, cnidarians, and bilaterians. The "phylogenomic" approach [1] has recently provided a robust picture for intrabilaterian relationships [2, 3] but not yet for more early branching metazoan clades. We have assembled a comprehensive 128 gene data set including newly generated sequence data from ctenophores, cnidarians, and all four main sponge groups. The resulting phylogeny yields two significant conclusions reviving old views that have been challenged in the molecular era: (1) that the sponges (Porifera) are monophyletic and not paraphyletic as repeatedly proposed [4-9], thus undermining the idea that ancestral metazoans had a sponge-like body plan; (2) that the most likely position for the ctenophores is together with the cnidarians in a "coelenterate" clade. The Porifera and the Placozoa branch basally with respect to a moderately supported "eumetazoan" clade containing the three taxa with nervous system and muscle cells (Cnidaria, Ctenophora, and Bilateria). This new phylogeny provides a stimulating framework for exploring the important changes that shaped the body plans of the early diverging phyla.
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                Author and article information

                Journal
                Mol Biol Evol
                molbiolevol
                molbev
                Molecular Biology and Evolution
                Oxford University Press
                0737-4038
                1537-1719
                September 2010
                08 April 2010
                08 April 2010
                : 27
                : 9
                : 1983-1987
                Affiliations
                [1 ]Department of Earth and Environmental Sciences, Palaeontology and Geobiology GeoBio-Center LMU, Ludwig-Maximilians-Universität München, München, Germany
                [2 ]Department of Geobiology, Courant Research Center Geobiology, Georg-August Universität Göttingen, Göttingen, Germany
                [3 ]Centre Robert-Cedergren, Département de Biochimie, Université de Montréal, Montréal, Québec, Canada
                [4 ]Abteilung Bioinformatik, Institut für Mikrobiologie und Genetik, Georg-August-Universität Göttingen, Göttingen, Germany
                [5 ]Department of Applied Molecular Biology, Institute for Physiological Chemistry and Pathobiochemistry, Mainz, Germany
                [6 ]University Pierre & Marie Curie (UPMC), Centre National de la Recherche Scientifique, Muséum National d'Histoire Naturelle, Department UMR7138 Systématique, Adaptation, Evolution. UPMC, Paris, France
                Author notes
                [†]

                These authors contributed equally to the present study.

                [* ] Corresponding author: E-mail: woerheide@ 123456lmu.de .

                Associate editor: Manolo Gouy

                Article
                10.1093/molbev/msq089
                2922619
                20378579
                d6949997-276d-410b-bcc9-4a95c110a629
                © The Author(s) 2010. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.

                This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License ( http://creativecommons.org/licenses/by-nc/2.5), which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

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                Letters

                Molecular biology
                long-branch attraction,saturation,metazoa,ctenophora,est,multigene analysis,porifera
                Molecular biology
                long-branch attraction, saturation, metazoa, ctenophora, est, multigene analysis, porifera

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