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Australian Beetles Volume 2: Archostemata, Myxophaga, Adephaga, Polyphaga (part) PDF

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Preview Australian Beetles Volume 2: Archostemata, Myxophaga, Adephaga, Polyphaga (part)

Volume 2 Volume 2 This three-volume series represents a comprehensive treatment of the A beetles of Australia, a relatively under-studied fauna that includes u many unusual and unique lineages found nowhere else on Earth. s t Volume 2 contains 36 chapters, providing critical information r and identification keys to the genera of the Australian beetle families a included in suborders Archostemata, Myxophaga, Adephaga and several l i groups of Polyphaga (Scirtoidea, Hydrophiloidea, Scarabaeoidea, Buprestoidea a and Tenebrionidae). Each chapter is richly illustrated in black and white drawings n and photographs. The book also includes colour habitus figures for about 1000 B Australian beetle genera and subgenera belonging to the families treated in this volume. e This volume is a truly international collaborative effort, as the chapters have been written by 23 e AusAtursatrlailaiann B Beeetleestles contributors from Australia, China, Czech Republic, Germany, Italy, Poland and USA. t l e Morphology, Classification and Keys About the authors s Archostemata, Myxophaga, Adephaga, Polyphaga (part) Adam Ślipiński completed his PhD and DSc in Poland, where he worked for 20 years at the Museum and Institute of Zoology of the Polish Academy of Sciences, Warsaw; he also held a E A joint appointment as Professor of Biology at the University of Zielona Gora. He is currently d r i c working as a Senior Principal Research Scientist and curator at the Australian National to h r o s s Insect Collection, CSIRO. He is the author of nearly 200 research publications, the editor of : t A e m a two-volume book on the phylogeny and classification of beetles, and an author of books on d a a Australian ladybird and longhorn beetles. Adam’s research concentrates on the phylogeny and m ta , higher classification of beetles. Ś M l ip y i x John F. Lawrence is one of the world’s preeminent coleopterists. He worked at Harvard ń o s p University’s Museum of Comparative Zoology before emigrating to Australia, where he served k h i a as a Senior Principal Research Scientist at the Australian National Insect Collection, CSIRO, a g n a until 1999. He has published more than 200 research articles and book chapters and was d , A J co-editor of a recent volume of Handbook of Zoology, in which he authored or co-authored more o d h e than 50 chapters. John is now retired but continues to conduct research on the phylogeny and n p h F classification of beetles. . a L g a a , w P r o e l n y c p e h a g a ( p a r t ) Editors: Adam Ślipiński and John F. Lawrence Volume 2 Australian Beetles Archostemata, Myxophaga, Adephaga, Polyphaga (part) Editors: Adam Ślipiński and John F. Lawrence Copyright The Authors 2019. All rights reserved. Except under the conditions described in the Australian Copyright Act 1968 and subsequent amendments, no part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, duplicating or otherwise, without the prior permission of the copyright owner. Contact CSIRO Publishing for all permission requests. The authors and editors each assert their moral rights, including the right to be identified as the author or editor. A catalogue record for this book is available from the National Library of Australia. Published by: CSIRO Publishing Locked Bag 10 Clayton South VIC 3169 Australia Telephone: +61 3 9545 8400 Email: [email protected] Website: www.publish.csiro.au Cover images by Cate Lemann and Adam Ślipiński Cover design by Andrew Weatherill Typeset by Thomson Digital Printed in China by Leo Paper Products Ltd. CSIRO Publishing publishes and distributes scientific, technical and health science books, magazines and journals from Australia to a worldwide audience and conducts these activities autonomously from the research activities of the Commonwealth Scientific and Industrial Research Organisation (CSIRO). The views expressed in this publication are those of the author(s) and do not necessarily represent those of, and should not be attributed to, the publisher or CSIRO. The copyright owner shall not be liable for technical or other errors or omissions contained herein. The reader/user accepts all risks and responsibility for losses, damages, costs and other consequences resulting directly or indirectly from using this information. The paper this book is printed on is in accordance with the standards of the Forest Stewardship Council® and other controlled material. The FSC® promotes environmentally responsible, socially beneficial and economically viable management of the world’s forests. Jul19_01 CONTENTS PREFACE � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � v DEDICATION� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � vi ACKNOWLEDGMENTS� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � vi CONTRIBUTORS TO VOLUME 2 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � vii 1� Archostemata Kolbe, 1908� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 1 John F. Lawrence and Hermes E. Escalona 2� Ommatidae Sharp and Muir, 1912� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 4 John F. Lawrence and Hermes E. Escalona 3� Cupedidae Laporte, 1836 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 8 John F. Lawrence and Hermes E. Escalona 4� Myxophaga Crowson, 1955� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 13 John F. Lawrence 5� Sphaeriusidae Erichson, 1845� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 15 W. Eugene Hall 6� Adephaga Schellenberg, 1806� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 18 John F. Lawrence 7� Gyrinidae Latreille, 1810� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 20 Chris H.S. Watts 8� Haliplidae Aube, 1836 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 24 Chris H.S. Watts 9� Noteridae Thomson, 1860 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 27 Tom A. Weir and Cate Lemann 10� Hygrobiidae Régimbart, 1879� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 32 Lars Hendrich 11� Dytiscidae Leach, 1815� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 34 Lars Hendrich, Cate Lemann and Tom A. Weir 12� Carabidae Latreille, 1802 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 61 Martin Baehr and Kipling Will 13� Polyphaga Emery, 1886� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 218 John F. Lawrence 14� Superfamily Scirtoidea Fleming, 1821� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 219 John F. Lawrence 15� Scirtidae Fleming, 1821� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 221 Chris H.S. Watts and Peter Zwick 16� Superfamily Hydrophiloidea Leach, 1815� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 249 Martin Fikáˇcek 17� Georissidae Castelnau, 1840� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 251 Martin Fikáˇcek 18� Hydrochidae Thomson, 1859 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 258 Martin Fikáˇcek 19� Spercheidae Erichson, 1837 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 265 Martin Fikáˇcek 20� Hydrophilidae Leach, 1815� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 271 Martin Fikáˇcek 21� Histeridae Gyllenhal, 1808 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 338 Yu-Lingzi Zhou, Sławomir Mazur, Tomáš Lackner and Adam Ślipiński iv Australian Beetles 22� Superfamily Scarabaeoidea Latreille, 1802� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 362 John F. Lawrence, Nicole L. Gunter and Tom A. Weir 23� Geotrupidae Latreille, 1802 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 369 John F. Lawrence and Mengjie Jin 24� Passalidae Leach, 1815 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 377 Eric G. Matthews and Owen D. Seeman 25� Trogidae Macleay, 1819� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 387 John F. Lawrence 26� Lucanidae Latreille, 1804 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 391 Chris Reid 27� Hybosoridae Erichson, 1847� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 417 Eric G. Matthews and Alberto Ballerio 28� Scarabaeidae Latreille, 1802� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 425 John F. Lawrence and Tom A. Weir 29� Scarabaeidae: Aphodiinae Leach, 1815� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 431 Tom A. Weir, John F. Lawrence, Cate Lemann and Nicole L. Gunter 30� Scarabaeidae: Scarabaeinae Latreille, 1802 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 443 Nicole L. Gunter, Cate Lemann and Tom A. Weir 31� Scarabaeidae: Melolonthinae Leach, 1819 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 467 Tom A. Weir, John F. Lawrence, Cate Lemann and Nicole L. Gunter 32� Scarabaeidae: Rutelinae Macleay, 1919 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 508 Tom A. Weir, John F. Lawrence, Cate Lemann and Nicole L. Gunter 33� Scarabaeidae: Dynastinae Macleay, 1919 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 516 Tom A. Weir, John F. Lawrence, Cate Lemann and Nicole L. Gunter 34� Scarabaeidae: Cetoniinae Leach, 1815 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 531 Christian H. Moeseneder, Tom A. Weir, Cate Lemann and Paul M. Hutchinson 35� Buprestidae Leach, 1815� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 554 John F. Lawrence and Cate Lemann 36� Tenebrionidae Latreille, 1802� � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 582 Eric G. Matthews and John F. Lawrence COLOUR PLATES � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 663 ERRATA FOR VOLUME 1 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 766 INDEX OF GENERIC AND FAMILY GROUP NAMES � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 769 PREFACE As indicated in the preface to Volume 1 of this series, the chapters with one major exception. Hydrophilidae is now broken up into in the present volume and those following it, have been or will several families, of which the following four have Australian be written by specialists from various parts of the world, as well representatives: Georissidae, Hydrochidae, Spercheidae and as by the two editors and other members of the CSIRO staff and Hydrophilidae (including Sphaeridiinae); this is not a new students. Although the original plan was to include in this volume concept and the change probably should have been made in the Australian families of Archostemata, Myxophaga, Adephaga, previous volume. Hydrophiloidea, Staphylinoidea and Scarabaeoidea, the current Below the level of family, the changes are often more availability of family specialists necessitated some reorganisation. considerable. The classification of Carabidae in Volume 1, for As a result, Staphylinoidea will now be included in Volume 3, instance, included 13 Australian subfamilies, with most tribes the families Scirtidae, Buprestidae and Tenebrionidae have been included in Harpalinae. In Chapter 12 of the present volume, no moved to the present volume, and chapter numbers will not be subfamilies are recognised, and genera are placed in 37 tribes and consistent with those used in Volume 1. numerous subtribes. Changes in subfamilial, tribal and generic It is also well known that the family level classification of concepts have occurred in various other families based on newly Coleoptera has changed considerably from that discussed and published data, and these are reflected in several chapters. outlined in Volume 1, Chapter 2. This is due in part to several In addition to the Staphylinoidea and the remaining Polyphaga worldwide, in-depth studies, involving both adult and larval the Volume 3 will include new sets of family level keys to adult morphology, and in part to the increase in molecular sequence and larvae reflecting ongoing changes in the higher classification data and improvements in phylogenetic analysis. In the present of beetles over the years. volume, most Australian family limits have remained the same, DEDICATION This volume is dedicated to the memory of A return visit to Queensland in 1981 allowed him to concentrate on carabid collecting and turned his interest to the Australian MARTIN BAEHR fauna. He returned for extended field and museum trips every two years for the rest of his life, exploring every corner of the March 10, 1943 – April 17, 2019 continent and often spending weeks alone in remote areas like the Kimberley, Kakadu and Cape York. His prodigious collections Dr Martin Baehr served his career as a curator at the Bavarian were matched by his research output and he published 187 State Collection of Zoology (Zoologische Staatssammlung papers totalling 5789 pages on the Australian biota and described München) in Munich, Germany as a leading authority on 1167 new Carabidae from Australia as well as 52 Araneae, 13 the Carabidae of the Indo-Australian region. He first visited Hemiptera, 1 Onychophora and 33 Orthoptera. In parallel, he Australia and Tasmania for a year in 1972–73, and fell in love published another 183 papers on non-Australian Carabidae, with the country, though then being interested in Onychophora, mostly from SE Asia and the islands through to New Guinea. reptiles and arachnids. His PhD studies at Tübingen were on the His final, and most comprehensive, work is the generic overview functional structure of the thorax and legs of Carabidae and he and synthesis of the whole Australian carabid fauna, written in began to work on their taxonomy, first on the European fauna, collaboration with Kipling Will, which appears in this volume, then a short 1976 paper on the Carabidae from his Tasmanian visit. and which he sadly did not live to see in print. ACKNOWLEDGMENTS We gratefully acknowledge The Australian National Insect and graphics skills, and Cate Lemann for photography, testing the Collection, CSIRO for support of research and for allowing keys and continuous technical assistance. the use of copyrighted images. We are also in debt to the many We thank the following photographers for providing photos institutions and individuals who have supported our various of living beetles: Kristi Ellingsen, Simon Grove, Owen Kelly, research projects over the years by supplying specimens, literature Zhenhua Liu, Jiří and Marie Lochman (Lochman Transparencies) and information. Anne Hastings is acknowledged for her artwork and Nick Monaghan (Life Unseen). CONTRIBUTORS TO VOLUME 2 Editors: Adam Ślipiński and John F� Lawrence Australian National Insect Collection CSIRO GPO Box 1700 Canberra, ACT 2601, Australia Martin Baehr GPO Box 1700 Zoologische Staatssammlung Canberra, ACT 2601, Australia Münchhausenstraße 21 Eric G� Matthews 81247 München, Germany South Australian Museum Alberto Ballerio North Terrace Viale Venezia 45 Adelaide, SA 5000, Australia I-25123 Brescia, Italy Sławomir Mazur Hermes E� Escalona Department of Forest Protection and Ecology Zoologisches Forschungsmuseum Alexander Koenig Warsaw University of Life Sciences Centre for Molecular Biodiversity Research Nowoursynowska 159 Adenauerallee 160 02–776 Warszawa, Poland 53113 Bonn, Germany Chris H� Moeseneder Martin Fikáček Oceans and Atmosphere Flagship, CSIRO Department of Entomology Queensland Biosciences Precinct National Museum Natural History 306 Carmody Road Cirkusová 1740 CZ-193 00 Praha 9 - St. Lucia, Qld 4067, Australia Horní Počernice, Czech Republic Chris A�M� Reid Nicole L� Gunter Australian Museum Department of Invertebrate Zoology 1 William Street Cleveland Museum of Natural History Sydney, NSW 2010, Australia Cleveland, Ohio 44106, USA Owen D� Seeman W� Eugene Hall Queensland Museum University of Arizona Insect Collection PO Box 3300 Department of Entomology South Brisbane, Qld 4101, Australia 1140 E. South Campus Dr Chris H�S� Watts Tucson, Arizona 85721, USA South Australian Museum Lars Hendrich North Terrace Zoologische Staatssammlung Adelaide, SA 5000, Australia Münchhausenstraße 21 Tom A� Weir 81247 München, Germany Australian National Insect Collection Paul M� Hutchinson CSIRO Quarantine WA GPO Box 1700 Department of Primary Industries and Regional Development, Canberra, ACT 2601, Australia Level 1, 24 Fricker Road Kipling W� Will Perth Airport, WA 6105, Australia Essig Museum of Entomology Mengjie Jin University of California, Berkeley School of Life Sciences Berkeley, California 94720, USA Sun Yat-Sen University Yu-Lingzi Zhou Guangzhou 510275, China Key Laboratory of Zoological Systematics and Evolution Tomáš Lackner Institute of Zoology, Chinese Academy of Sciences Zoologische Staatssammlung 1 Beichen West Road Münchhausenstraße 21 Beijing 100101, China 81247 München, Germany Peter Zwick Cate Lemann Schwarzer Stock 9 Australian National Insect Collection D-36110 Schlitz, Germany CSIRO This page intentionally left blank 1. ARCHOSTEMATA KOLBE, 1908 John F. Lawrence and Hermes E. Escalona The genera Cupes Latreille and Omma Newman were included families Ommatidae and Cupedidae. Recent discoveries of in the suborder Adephaga by Ganglbauer (1903), Kolbe (1901, Cupedidae in the Cretaceous amber of Myanmar may be found 1903) and Lameere (1903), based primarily on wing venation, in the papers by Jarzembowski et al. (2017a, 2017b, 2017c). while Kolbe (1908) excluded cupedids from Adephaga Some important characters defining this suborder may be and placed them in a separate group, Archostemata, within found in extant members of the Ommatidae and Cupedidae, but what is now known as Polyphaga. Forbes (1926) considered not in highly derived or little known groups like Micromalthidae Archostemata to be one of the three suborders of beetles, (Beutel & Hörnschemeyer 2002a), Crowsoniellidae (Ge et al. containing Cupedidae Laporte (including Ommatidae Sharp 2011) or the doubtfully included Jurodidae (Yan et al. 2014). & Muir) and Micromalthidae Barber, based on wing folding, These include: (1) lack of cervical sclerites (also in Adephaga while by Böving & Craighead (1931) and Peyerimhoff (1933) and Myxophaga); (2) external propleuron extending well in front supported this based on larval evidence. This classification was of procoxal cavities (also in Adephaga, reduced in Myxophaga); used in Crowson’s (1955) classic work on the order. In a later (3) free, external protrochantin; (4) membranous joint between paper, Crowson (1962) made the first attempt to integrate the mesothorax and metathorax (also in some Polyphaga); (5) work of paleontologists, such as A. Handlirsch, R. J. Tillyard, mesocoxal cavities broadly closed laterally by metanepisternum B. B. Rohdendorf and A. B. Martynov, on cupedid-like fossils (narrowly so in a few Polyphaga); (6) well developed and with knowledge of the extant archostematan fauna known at exposed metatrochantins; (7) hind wing without a radial cell that time. Ponomarenko (1964, 1966, 1968, 1969) published formed by meeting of RA and RA (also in Adephaga 1+2 3+4 the first major work dealing with Archostemata, which was and Myxophaga); (8) hind wing with cross-veins between RP defined in a broad sense to include virtually all known Permian and MP forming an oblongum cell (also in Adephaga and 1+2 and Mesozoic fossils sufficiently complete to be described (11 Myxophaga), (9) hind wing with major fold crossing MP 1+2 families, 74 genera and 186 species) but lacking the distinctive forming a sharp hinge (also in Adephaga and Myxophaga); (10) features of either Polyphaga or Adephaga. The Lower Permian larva with 6-segmented legs and paired pretarsal claws (also in Tshekardocoleidae and several Upper Permian groups were Adephaga); (11) larva with ligular sclerome. placed in Protocoleoptera and Archecoleoptera, respectively, There are relatively few extant species of Archostemata: by Crowson (1975), but were considered to form a coleopterid the family Ommatidae contains four Australian species of stem group by Kukalová-Peck (1991). This was followed by Omma Newman and two species of Tetraphalerus Waterhouse, Beutel (1997) and Beutel & Friedrich (2008), who added the occurring primarily in Argentina and Bolivia, and the Cupedidae Triassic Triadocupedidae to this paraphyletic stem group. includes nine genera and 31 species occurring in North America, Within the remaining Archostemata, relationships are still South America, the eastern Palaearctric and Oriental regions, not clearly understood, particularly among the taxa with non- the East Indies, New Guinea, Australia, New Caledonia, East clathrate elytra (without window-punctures) such as Jurodidae, Africa, South Africa and Madagascar (Neboiss 1984; Lawrence Ademosynidae, Schizophoridae and Catiniidae, and their 1999; Hörnschemeyer 2009, 2016; Hörnschemeyer & Beutel possible affinities with the other three suborders, Adephaga, 2016; Hörnschemeyer & Yavorskaya 2016). The group has Myxophaga and Polyphaga. Numerous publications have become extinct in Europe, but several European species are appeared in the past 25 years on fossil Archostemata primarily known from the early Tertiary (Kirejtshuk 2005; Kirejtshuk et from localities in Kazakhstan, Russia, Mongolia and northern al. 2010a). Several morphological papers have been produced China but also on those in the UK, USA, Spain, Australia and on both larvae (Beutel & Hörnschemeyer 2002a, 2002b; Argentina; included are Ashman et al. (2015); Jarzembowski & Hörnschemeyer et al. 2002; Grebennikov 2004; Yavorskaya et Wang (2016); Jarzembowski et al. (2012, 2013a, 2013b, 2015); al. 2015) and adults (Baehr 1975; Hörnschemeyer et al. 2006; Kirejtshuk (2005); Kirejtshuk et al. (2010a, 2010b, 2016); Beutel et al. 2008; Friedrich et al. 2009; Hünefeld et al. 2011). Lubkin (2003); Martin (2010); Martins-Neto et al. (2006); General summaries of extant Archostemata may be found in Ponomarenko (1994, 1997, 2000, 2004, 2006); Ponomarenko Hörnschemeyer (2016), Hörnschemeyer & Beutel (2016) and & Martinez-Delclòs (2000); Ponomarenko & Ren (2010); Hörnschemeyer & Yavorskaya (2016). Ponomarenko et al. (2014); Ren & Tan (2006); Ren et al. (2006); Soriano & Delclòs (2006); Tan & Ren (2006a, 2006b, References 2006c, 2007, 2009); Tan et al. (2007a); Tan et al. (2005); Tan Ashman LG, Oberprieler RG, Ślipiński A (2015) Rhopalomma stefaniae et al. (2006a, 2006b, 2007b, 2007c); Tan et al. (2006c, 2013, gen. et sp. n., the first ommatid beetle from the Upper Jurassic in Aus- 2012) and Yan et al. (2014). tralia (Coleoptera: Archostemata: Ommatidae). Zootaxa 3980(1), 136– 142. doi:10.11646/zootaxa.3980.1.8 In the study of Archostemata by Kirejtshuk et al. (2016), Baehr M (1975) Skelett und Muskulatur des Thorax von Priacma serrata virtually all archostematans with clathrate or reticulate elytra are LeConte (Coleoptera, Cupedidae). Zeitschrift für Morphologie der Tiere considered to belong to the single family Cupedidae, extending 81, 55–101. doi:10.1007/BF00290073 back to the Triassic, but without an adequate phylogenetic Beutel RG (1997) Über Phylogenese und Evolution der Coleoptera (In- analysis and additional support from other members of the secta), insbesondere der Adephaga. Abhandlungen des Naturwissen- paleontological community, we continue to recognise the schaftlichen Vereins in Hamburg (NF) 31, 164 pp.

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