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Functional anatomy of the explosive defensive system of bombardier beetles (Coleoptera,Carabidae, Brachininae)
Affiliation:1. Department of Science, University Roma Tre, Viale G. Marconi, 446, 00146 Roma, Italy;2. Department of Agricultural, Food and Environmental Sciences, University of Perugia, Borgo XX Giugno 74, 06121 Perugia, Italy;1. Department of Ethology and Social Biology of Insects, Schmalhausen-Institute of Zoology, B. Chmielnicki Street 15, 01601 Kiev, Ukraine;2. Department of Functional Morphology and Biomechanics, Christian-Albrechts-University of Kiel, Am Botanischen Garten 1-9, D-24098 Kiel, Germany;3. Institute for Evolutionary Ecology, Acad. Lebedev Street 37, 03143 Kiev, Ukraine;4. Department of Entomology, Biological Faculty, Lomonosov Moscow State University, 1-12 Leninskie Gory, 119991 Moscow, Russia;5. Department of Scientific Fundamental Collection, Schmalhausen-Institute of Zoology, B. Chmielnicki Street 15, 01601 Kiev, Ukraine;6. Department of Zoology, Dragomanov Pedagogical University, Pirogov Street 9, 01601 Kiev, Ukraine;7. Department of Zoology, Institute of Biology, National Shevchenko-University, Volodymirska Street, 64, 01601 Kiev, Ukraine;1. Institute of Biology, Faculty of Natural Sciences, Martin Luther University Halle-Wittenberg, Heinrich-Damerow Str. 4, 06120 Halle, Germany;1. Center of Studies on Social Insects, Bioscience Institute, UNESP – Universidade Estadual Paulista, Campus Rio Claro-SP, Brazil;2. Biology Department, Biosciences Institute, UNESP – Universidade Estadual Paulista, Campus Rio Claro-SP, Brazil;3. Behavioral Physiology and Sociobiology (Zoology II), Biocenter, University of Würzburg, Germany;1. Czech University of Life Sciences, Faculty of Forestry and Wood Sciences, Prague, Czech Republic;2. School of Biological Sciences, University of Sydney, Sydney, NSW, Australia;3. Museu de Zoologia da Universidade de São Paulo, 04218-970, Av. Nazaré, 481, Ipiranga, São Paulo, CEP 04263-000, São Paulo, SP, Brazil;4. Centre des Sciences du Goût et de l’Alimentation, UMR 6265 CNRS, UMR 1324 INRA, Université de Bourgogne, Agrosup Dijon, 6 Bvd Gabriel, 21000, Dijon, France;1. Instituto de Ciencias Naturales, Universidad Nacional de Colombia, A.A. 7495, Bogotá, Colombia;2. Zoological Institute, K.U. Leuven, Naamsestraat 59, Box 2466, B-3000, Belgium
Abstract:This paper provides the first comparative anatomical study of the explosive pygidial defensive system of bombardier beetles in species classified in three brachinine subtribes: Brachinus (Brachinina), Pheropsophus (Pheropsophina) and Aptinus (Aptinina). We investigated the morphology and ultrastructure of this system using optical, fluorescence, and focused ion beam (FIB/SEM) microscopy. In doing so, we characterized and comparatively discussed: (1) the ultrastructure of the gland tissues producing hydroquinones and hydrogen peroxide (secretory lobes), and those producing catalases and peroxidases (accessory glands); (2) the complex anatomy of the collecting duct; (3) the arrangement of the muscular bundles and the folding of the cuticle of the reservoir, suggesting a functional division of this chamber (dynamic part and storage part); (4) the great structural diversity of sculpticles inside the reaction chamber, where we could recognize six main types of microsculpture located in specific districts of the chamber. Additionally, using fluorescence microscopy, we highlighted the presence of resilin in two structures strongly subjected to mechanical stress during the discharge, the valve and the turrets of the reaction chamber. The results of this paper give a solid anatomic overview of the most popular beetle defensive system, contributing to the debate on its evolution within the Carabidae.
Keywords:Bombardier beetles  Brachininae  Explosive glands  FIB/SEM  Ultrastructure  Pygidial glands  Carabidae
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