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The gonangulum: A reassessment of its morphology,homology, and phylogenetic significance
Authors:Klaus-Dieter Klass  Natalia A Matushkina  Johanna Kaidel
Institution:1. Senckenberg Natural History Collections Dresden, Museum of Zoology, Königsbrücker Landstraße 159, 01109 Dresden, Germany;2. Department of Zoology, Biological Faculty, National Taras Shevchenko University of Kyiv, vul. Volodymirs''ka 64, 01601 Kyiv, Ukraine;1. Institut de Systématique, Évolution, Biodiversité (ISYEB) Muséum national d''Histoire naturelle, CNRS, Sorbonne Université, EPHE, Université des Antilles, CP50, 57 rue Cuvier, 75005, Paris, France;2. Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY, 10024, USA;3. Division of Entomology, Natural History Museum, and Department of Ecology & Evolutionary Biology, University of Kansas, 1501 Crestline Drive – Suite 140, Lawrence, KS, 66045, USA;1. State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 East Beijing Road, Nanjing 210008, China;2. Department of Earth Sciences, The University of Hong Kong, Hong Kong Special Administrative Region, China;3. Institut de Systématique, Évolution, Biodiversité, ISYEB-UMR 7205-CNRS, MNHN, UPMC, EPHE, Muséum National d''Histoire Naturelle, Sorbonne Universités, 57 rue Cuvier, CP 50, Entomologie, F-75005, Paris, France;4. Department of Earth Sciences, The Natural History Museum, London SW7 5BD, UK;5. Goglio (Tianjin) Packaging Co. Ltd, No. 99, West 11th Avenue, Airport Economic Area, Tianjin, China;6. Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, 1, Beichen West Road, Beijing 100101, China;1. Systematic Entomology, Graduate School of Agriculture, Hokkaido University, Sapporo 060-8589, Japan;2. Muséum d’histoire naturelle, C.P. 6434, CH-1211 Genève 6, Switzerland;1. Institut de Systématique, Évolution, Biodiversité (ISYEB), Muséum national d’Histoire naturelle, CNRS, Sorbonne Université, EPHE, Université des Antilles, CP50, 57 rue Cuvier 75005 Paris, France;2. CNRS, UMR 5554 Institut des Sciences de l’Évolution de Montpellier, Place Eugène Bataillon, 34095, Montpellier, France;3. Univ. Rennes, CNRS, Géosciences Rennes, UMR 6118, F-35000, Rennes, France;4. 178 rue des Plattières, F-38300 Nivolas-Vermelle, France;1. Ludwig-Maximilians-Universität München (LMU Munich), Biocenter, Großhaderner Str. 2, 82152 Planegg-Martinsried, Germany;2. GeoBio-Center at LMU, Richard-Wagner-Str. 10, 80333 München, Germany;3. Kreuzbergstr. 90, 66482 Zweibrücken, Germany;1. Department of Life Sciences, University of Siena, Via A. Moro 2, I-53100 Siena, Italy;2. Sugadaira Montane Research Center, University of Tsukuba, Nagano 386-2204, Japan
Abstract:The gonangulum is a sclerite in the female genitalic region of insects. Its presence or full development has long been considered an apomorphy supporting Zygentoma + Pterygota. Recent studies of female genitalia in several insect orders (K.-D. Klass and co-workers) revealed many new data on the gonangulum and homologous sclerotisations (laterocoxa LC9). Herein the gonangulum area is described (including articulations, muscle attachments, sulci) and compared among Archaeognatha, Zygentoma, Odonata, Dermaptera, Dictyoptera, and Notoptera. A wider perspective is provided to the topic by addressing some novel issues: identification of LC9 sclerotisations in non-insect taxa and in insects that secondarily lack an ovipositor; occurrence of homonomous sclerotisations in other abdominal segments of both sexes; morphological interpretation of LC9; and the role of paedomorphosis in LC9 evolution. As a result, there is currently no support for any insect lineage from this character system. For gonangulum-related characters both a significant intra-ordinal variation and frequent homoplasy are demonstrated using various Odonata, Dermaptera, and Dictyoptera as examples. Divergent fates of LC9 in simplified genitalia are shown using a dermapteran and an odonatan. We view all this as a showcase of how a renewed and more detailed examination of a character system can dramatically change the phylogenetic evidence drawn from it.
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