Handbook of Zoology / Handbuch der Zoologie: Teilband/Part 38 Volume 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim) 9783110904550, 9783110171303

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Handbuch der Zoologie Handbook of Zoology Band/Volume IV Arthropoda: Insecta Herausgeber/Editor Niels P. Kristensen & Rolf G. Beutel Coleoptera, Beetles Volume 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim) Teilband/Part 38

Handbuch der Zoologie Eine Naturgeschichte der Stämme des Tierreiches

Handbook of Zoology Α Natural History of the Phyla of the Animal Kingdom

Gegründet von / Founded by Willy Kükenthal Fortgeführt von / Continued by Μ. Beier, Μ. Fischer, J.-G. Helmcke, D. Starck, H. Wermuth

Band / Volume IV Arthropoda: Insecta

Teilband / Part 38

Herausgeber / Editors Niels P. Kristensen & Rolf G. Beutel

W DE G

Walter de Gruyter · Berlin · New York

Rolf G. Beutel Richard Α. B. Leschen (Volume Editors)

Coleoptera, Beetles Volume 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim)

W DE G Walter de Gruyter · Berlin · New York

Herausgeber Scientific Editors Professor Dr. Rolf G. Beutel Institut für Spezielle Zoologie und Evolutionsbiologie Friedrich-Schiller-Universität Jena 07743 Jena Germany

Professor Dr. Niels P. Kristensen Zoological Museum University of Copenhagen Universitetsparken 15 DK-2100 Copenhagen Denmark

Verlag / Publishers Walter de Gruyter G m b H & Co. K G Genthiner Straße 13 10785 Berlin Germany

Library of Congress — Cataloging-in-Publication

Walter de Gruyter, Inc. 500 Executive Boulevard Ossining, N Y 10562 USA

Data

Handbook of Zoology

agr 23001436

Front cover image: The cover shows ground beetles by F.Cassola (1976), S. W. Nichols (1985) und G.S.Lafer (1989). ® Printed on acid-free paper which falls within the guidelines of the ANSI to ensure permanence and durability.

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ISBN 3-11-017130-9

Editors' preface

With approximately 350000 described species, Coleoptera are presently by far the most speciesrich order of insects and largest group of animals of comparable age. This is a well known fact and the resulting enormous ecological and economic implications are almost common-place today. As the knowledge about our world increases, the more important are general up-to-date works that cover natural history, systematics, and evolution of beetles. The immense biodiversity of Coleoptera is challenging and somewhat intimidating for biologists, and an attempt to treat three of four suborders and more than one third of the families (60) in one of three planned Coleoptera handbook volumes may seem almost presumptious to us. Nevertheless, we felt confident that a compilation of presently available data, including results of recent investigations, would be worth the effort, and would prove very useful for coleopterists and other biologists needing access to phylogenetic, taxonomic, morphological and natural history information. The first of three planned Coleoptera volumes covers the suborders Archostemata, Myxophaga and Adephagha, the series Staphyliniformia, Scarabaeiformia and Elateriformia (excluding Elateroidea) of Polyphaga, and some introductory chapters on the early evolution of beetles, morphology, and recent advances in molecular systematics. Initially we considered to include a chapter on the history of beetle classification. However, we finally decided against it, as this topic was treated comprehensively by J. F. Lawrence and A. F. Newton Jr. (1995) in the volumes dedicated to the 80 th birthday of Roy A. Crowson, and no major changes of classification were proposed since that time. The classification in the present volume is based on the family and subfamily treatment by these authors (with few modifications), which is presently widely accepted (though still in need of revision). We also decided not to include a key to families in this volume as interactive keys on compact discs and traditional keys in several other books are easily available. The second volume will cover the remaining polyphagan taxa and recently described groups, which could not be included in the present volume. The third planned volume will be dedicated to a more detailed treatment of different structural complexes (e.g., larval chaetotaxy and skeleto-muscular systems), organ systems (e.g., digestive tract and ovarioles), and evolution of beetles and their immature stages. Cross references between chapters in the Coleoptera Handbook will be treated like that

presented in the Lepidoptera volume and each chapter indicated by emboldened volume and chapter number: 1 — 10.2 and 2—10.2 refer to chapters 10.2 in volume 1 and 2, respectively. Earlier parts of the handbook series ("Handbuch der Zoologie") were published in German and were almost exclusively available to German speaking scientists. This policy was changed completely. Like the recently issued Lepidoptera volumes (Vol. IV, Arthropoda: Insecta, Lepidoptera, Moths and Butterflies I and II, N. P. Kristensen ed.), which were certainly milestones in the history of the series, this treatment of Coleoptera is the combined effort of an international team of authors, entirely published in English, and explicitly addressing the international scientific community. The purpose of this work is to present a comprehensive survey of the present state of systematics and comparative biology of the beetle families. In contrast to the new edition of the 'American Beetles', the handbook volumes cover the world Coleoptera, and included is information on the distribution of each family. Emphasis is placed on detailed morphological treatments and systematics, and anatomical data are presented for adults and immature stages whenever such information is available. Results of modern phylogenetic analyses are presented for taxa where the data exist. Though we recognise that few molecular studies have been published yet, these too have been consulted where appropriate. The format of the chapters is largely consistent. However, for pragmatic reasons we have not listed the references separately for the families in some cases. They are inserted at the end of the superordinate chapters on Myxophaga, Archostemata and Scarabaeoidea. The present knowledge and presentation for the taxa treated differs widely due to the relative amount of available information and to the extent of the chapter produced by the contributing authors. Some of the family-group taxa have been described only very recently and others are largely unknown in many respects due to their cryptic habits (e.g., Crowsoniella) or their occurence in very remote and inaccessible areas (e.g., Jurodidae). Meanwhile systematic studies of many groups is at their infancy, and in these cases it appeared useful to present recent stages of the taxonomic development, which is tremendous in terms of newly discovered or erected taxa in some groups such as Elmidae. Introductory chapters with emphasis on phylogeny are presented for the suborders, series,

VI and for the well established larger superfamilies. Interrelationships within the small superfamilies (Scirtoidea, 1 — 15, Dascilloidea, 1 — 16) are discussed in the family chapters and Byrrhoidea (1—18) is not discussed as a whole, as the concept of this large taxon is not yet well supported by phylogenetic investigations. A considerable number of prominent coleopterists have agreed to cooperate in this project. However, we have also encouraged younger scientists involved with beetle systematics to contribute to this volume. So, the list of chapter authors reflects this rather asymmetrical group of specialists and the need for the mixed generations of scientists to interact and produce a volume of the highest quality of information. We are especially honoured by the cooperation of John F. Lawrence (CSIRO, Canberra) and Alfred F. Newton Jr. (Field Museum of Natural History, Chicago, [FMNH]) for their unending support and contribution to Coleoptera systematics. Some of the authors have agreed to join the project at a very late stage and under difficult conditions. It is greatly appreciated that some excellent contributions have been arranged within a very short time. The project was supported by scientists (Ronald Bellstedt, Vasily V. Grebennikov, Ignacio Ribera) and institutions through the loan or gift of valuable material (Australian National Insect Collection, CSIRO; Deutsches Entomolgisches Institut, Greifswald; F M N H ; Naturhistorisches Museum Wien; New Zealand Arthropod Collection, Auckland; Zoologisk Museum, Copenhagen). This is gratefully acknowledged. Our special thanks go to Margret Roser (Institut für Spezielle Zoologie und Evolutionsbiologie, FSU Jena) who has made a tremendous contribution in terms of graphical work. Finally we also want to express our gratitude to Mrs. Martina Bach, Dr. Christine Bowinkelmann, Dr. Stephanie Dawson, Mrs. Anita Eppelin and Dr. Annika Meyer (Walter de Gruyter, Biosciences Division) for their suport of this project.

Editors' preface

The selection of authors for a considerable number of the beetle families covered in this volume has been difficult because of the lack of specialists on certain groups and the recent deaths of some of our colleagues. An untimely event that has overshadowed the production of this volume was the completely unexpected death of the outstanding specialist of Hydrophiloidea, Michael Hansen (Zoologisk Museum, Copenhagen). His death was a serious shock and loss for the community of coleopterists and therefore we decided to dedicate this volume to our distinguished colleague. We should also mention, that Sebastian Endrödy-Younga (Transvaal Museum, Pretoria), who also passed away relatively recently, could have contributed significantly to this volume. The impact of systematists on biological inquiry is enormous and far-reaching and we hope that the death of Roy A. Crowson at the end of the last century does not mark the end of a strong tradition of morphology and phylogeny. However, the lack of taxonomic expertise and the emphasis on molecular biology and/or science for profit in some countries indicates a serious problem for systematics, and the need for beetle morphologists and taxonomists in particular. We are continually amazed by the paradox of the almost undisputed importance of biodiversity studies emphasized on higher political and administrative levels and the loss of systematic expertise for some important groups based on poorly planned executive decisions. We envision that this and the following volumes on Coleoptera systematics will demonstrate the continued need for taxonomic research and that systematics studies are not antiquated or anachronistic disciplines but are truly highly engaging, academic, and necessary for the whole of biology. Rolf G. Beutel Richard A. B. Leschen

Contents

Editors' preface

V

Contributors 1.

IX

Systematic position and early evolution Rolf G. Beutel

1

2.

Classification

3.

Molecular systematics of Coleoptera: what has been achieved so far? Alfried Vogler

17

Coleoptera, Morphology Rolf G. Beutel & John F. Lawrence

23

4.

5.

11

...

Archostemata Thomas Hörnschemeyer Introduction, Phylogeny

5.2. Cupedidae

32

5.3. Micromalthidae

34

5.4. Ommatidae

38

5.5. Jurodidae

40

Myxophaga Introduction, Phylogeny Rolf G. Beutel

43

6.1. Lepiceridae Roberto Arce-Perez, Jose Luis Navarrete-Heredia & Rolf G. Beutel...

45

6.2. Torridincolidae Rolf G. Beutel & Sergio A. Vanin . . . .

46

6.3. Sphaeriusidae Rolf G. Beutel & Roberto Arce-Perez . . 48 6.4. Hydroscaphidae Sergio A. Vanin, Rolf G. Beutel & Roberto Arce-Perez 7.

Adephaga Introduction, Phylogeny Rolf G. Beutel & Ignacio Ribera

49

119

7.9. Rhysodidae Rolf G. Beutel

145

8.

10.

Polyphaga Introduction, Phylogeny Rolf G. Beutel & Richard A. B. Leschen

153

Staphyliniformia Introduction, Phylogeny Rolf G. Beutel ά Richard A. B. Leschen

155

Hydrophiloidea Introduction, Phylogeny Miguel Archangelsky, Rolf G. Beutel & Albrecht Komarek

157

10.1. Hydrophilidae (Helophorinae, Epimetopinae, Georissinae, Hydrochinae, Spercheinae, Horelophinae, Horelophopsinae, Hydrophilinae, Sphaeridiinae) Miguel Archangelsky, Rolf G. Beutel ά Albrecht Komarek 10.2. Sphaeritidae Alfred Ν. Newton 10.3. Synteliidae Alfred N. Newton 10.4. Histeridae Peter W. Kovarik & Michael S. Caterino

53

64

7.3. Noteridae Konrad Dettner

72

7.4. Amphizoidae Konrad Dettner

81

7.5. Hygrobiidae Konrad Dettner

85

Staphylinoidea Introduction, Phylogeny Vastly V. Grebennikov 11.1. Hydraenidae Manfred Jäch, Rolf G. Beutel, Juan Antonio Delgado & Juan Angel Diaz 11.2. Ptiliidae W. Eugene Hall

158 183 187

190

11.

7.1. Gyrinidae Rolf G. Beutel & Robert E. Roughley . . 55 7.2. Haliplidae Bernard J. v. Vondel

115

7.8. Carabidae Erik Arndt, Rolf G. Beutel & Kipling Will

29 30

90

7.7. Trachypachidae Rolf G. Beutel & Erik Arndt

9.

5.1. Crowsoniellidae

6.

7.6. Dytiscidae Michael Balke

11.3. Agyrtidae Alfred F. Newton 11.4. Leiodidae Alfred F. Newton 11.5.

Scydmaenidae Sean O'Keefe

223

. 224 251 261 269 280

VIII 11.6. 11.7.

12.

Contents 15.3.

Silphidae Derek Sikes

288

Staphylinidae Margaret Thayer

296

15.4.

Scarabaeiformia Clarke Scholtz & Vastly V. Grebennikov Introduction

345

Definition

347

Families

349

Ancestral way of life

349

Evolutionary trends

354

Systematic position

355

Fossil record

16. 16.1. 16.2.

Clambidae Richard A. B. Leschen

438

Scirtidae John F. Lawrence

443

Dascilloidea Dascillidae John F. Lawrence

451

Rhipiceridae John F. Lawrence

456

17.

Buprestoidea Chuck Bellamy & Mark G. Volkovitsh 461

17.1.

Buprestidae

461

356

18.

Byrrhoidea

469

Larval morphology

357

18.1.

Byrrhidae Paul Johnson

469

13.

Scarabaeoidea

367

13.1.

Lucanidae

367

Elmidae Jan Kodada & Manfred Jäch

471

13.2.

Passalidae

369

13.3.

Trogidae

371

Dryopidae Jan Kodada & Manfred Jäch

496

13.4.

Glaresidae

373

18.4.

13.5.

Pleocomidae

373

Lutrochidae Cleide Costa, Sergio Vanin & Sergio Ide

508

13.6.

Bolboceratidae

374

18.5.

13.7.

Diphyllostomatidae

376

Limnichidae Ignacio Ribera & Carles Hernando . . 512

13.8.

Geotrupidae

377

18.6.

13.9.

Belohinidae

378

Heteroceridae Cleide Costa, Sergio Vanin, Sergio Ide & Rolf G. Beutel

518

18.7.

Psephenidae Chi-Feng Lee, Manfred Jäch & Rolf G. Beutel

521

18.8.

Cneoglossidae Cleide Costa, Sergio Vanin & Sergio Ide

533

Ptilodactylidae John F. Lawrence

536

13.10. Ochodaeidae

379

13.11. Ceratocanthidae

381

13.12. Hybosoridae

383

13.13. Glaphyridae

385

13.14. Scarabaeidae

386

14.

15. 15.1. 15.2.

Elateriformia (part.) Introduction, Phylogeny Richard A. B. Leschen & Rolf G. Beutel

18.2. 18.3.

18.9. 427

18.10. Chelonariidae Rolf G. Beutel & Richard A. B. Leschen

543 547 551

Scirtoidea Decliniidae John F. Lawrence

431

18.11. Eulichadidae Michael A. Ivie

Eucinetidae Richard A. B. Leschen

433

18.12. Callirhipidae John F. Lawrence

Contributors

Roberto Arce-Perez Departamento de Entomologia Instituto de Ecologia, A.C km. 2.5 antigua carretera a Coatepec No. 351, Congregation el Haya, C.P. 91070, Xalapa, Veracruz, Mexico e-mail: [email protected] Miguel Archangelsky Laboratorio de Ecologia Acuatica Universidad Nacional de La Patagonia Sarmiento 849 (9200) Esquel, Chubut, Argentina e-mail: [email protected] Erik Arndt Fachhochschule Anhalt Fachbereich LOEL Strenzfelder Allee 28, 06406 Bernburg, Germany e-mail: [email protected] Michael Balke Department of Entomology The Natural History Musum Cromwell Rd., London SW7 5BD, England e-mail: [email protected] Chuck Bellamy Plant Pest Diagnostics Branch California Department of Food & Agriculture 3294 Meadowview Road Sacramento, CA 95832, USA e-mail: [email protected] Rolf G. Beutel Institut für Spezielle Zoologie und Evolutionsbiologie FSU Jena 07743 Jena, Germany e-mail: [email protected] Michael S. Caterino Santa Barbara Museum of Natural History 2559 Puesta del Sol Road, Santa Barbara, CA 93105, USA e-mail: [email protected] Cleide Costa Museo de Zoologia Universidade de Säo Paulo Caixa Postal 42.694 04299-970 Säo Paulo, SP, Brasil e-mail: [email protected]

Juan Antonio Delgado Departamento de Biologia Animal (Zoologia) Facultad de Biologia Universidad de Murcia 30100 Murcia, Spain e-mail: [email protected] Konrad Dettner Lehrstuhl für Tierökologie II Universität Bayreuth Postfach 101251 95444 Bayreuth, Germany e-mail: [email protected] Juan Angel Diaz Departamento de Bioloxia Animal Facultade de Veterinaria Universidade de Santiago Campus de Lugo 27002 Lugo, Spain e-mail: [email protected] Vasily V. Grebennikov Department of Entomology University of Pretoria Pretoria 0002, South Africa e-mail: [email protected] W. Eugene Hall Zoology Section: Collections Manager CU Museum of Natural History UCB 265 University of Colorado Boulder, CO 80309-0265, USA e-mail: [email protected] Carles Hernando Museu de Ciencies Naturals de la Ciutadella Passeig Picasso s/n, Pare de la Ciutadella 08003 Barcelona, Spain e-mail: [email protected] Thomas Hörnschemeyer Institut für Zoologie und Anthropologie Abt. für Morphologie und Systematik Universität Göttingen Berliner Str. 28, 37073 Göttingen, Germany e-mail: [email protected] Sergio Ide Segäo de Entomologia Geral, Divisäo de Parasitologia Vegetal, Instituto Biologico, Caixa Postal 12.898 e 12.959, 04010-970 Säo Paulo SP, Brazil e-mail: [email protected]

χ Michael A. Ivie Department of Entomology Montana State University Bozeman, M T 59717, USA e-mail: [email protected] Manfred Jäch Naturhistorisches Museum 2. Zoologische Abteilung Burgring 7 A-1014 Wien, Österreich e-mail: manfred [email protected] Paul J. Johnson Insect Research Collection SD Natural History Collections and Biological Survey Box 2207A, South Dakota State University Brookings, SD 57007, USA e-mail: [email protected] Sean O'Keefe Department of Biological and Environmental Sciences Morehead State University Morehead, Kentucky 40351-1689, USA e-mail: [email protected] Jan Kodada Comenius University, Faculty of Natural Sciences Department of Zoology Mlynskä dolina B-l 84215 Bratislava, Slovakia e-mail: [email protected] Albrecht Komarek Naturhistorisches Museum 2. Zoologische Abteilung Burgring 7 A-1014 Wien, Österreich e-mail: [email protected] Peter W. Kovarik Division of Insects/Museum of Biodiversity 1315 Kinnear Rd. Columbus, OH 43212, USA e-mail: [email protected] John F. Lawrence 12 Hartwig Rd. Gympie, Queensland, 4570, Australia e-mail: [email protected] John. Lawrence@ento .csiro. au Chi-Feng Lee Institute of Zoology Academia Sinica Taipei, Taiwan 11529 e-mail: [email protected]

Contributors

Richard A. B. Leschen New Zealand Arthropod Collection Landcare Research, Private Bag 92-170 120 Mt Albert Road, Mt Albert Auckland, New Zealand e-mail: [email protected] Jose Luis Navarrete-Heredia Entomologia, Centro de Estudios en Zoologia Universidad de Guadalajara Apdo. Postal 234, 45100 Zapopan, Jalisco, Mexico e-mail: [email protected] Alfred F. Newton Division of Insects, Field Museum of Natural History Roosevelt Rd at Lake Shore Drive Chicago, Illinois 60605-2496, USA e-mail: [email protected] Ignacio Ribera Departamento de Biodiversidad y Biologia Evolutiva Museo Nacional de Ciencias Naturales Jose Gutierrez Abascal 2 28006 Madrid, Spain e-mail: [email protected] Robert E. Roughley Department of Entomology University of Manitoba Winnipeg, Manitoba Canada R 3 T 2 N 2 e-mail: [email protected] Clarke H. Scholtz Department of Entomology University of Pretoria Pretoria 0002, South Africa e-mail: [email protected] Derek S. Sikes Division of Zoology Department of Biological Sciences University of Calgary Calgary, Alberta, Canada, T2N 1N4 e-mail: [email protected] Margaret Thayer Div. of Insects, Field Museum of Natural History Roosevelt Rd at Lake Shore Drive Chicago, Illinois 60605-2496, USA e-mail: [email protected] Sergio Antonio Vanin Departamento de Zoologia Instituto de Biociencias Universidade de Säo Paulo

Contributors Caixa Postal 11461 05422-970 Säo Paulo, SP - Brasil e-mail: [email protected] Alfried P. Vogler The Natural History Museum Dept. of Entomology Cromwell Road London SW7 5BD, England e-mail: [email protected] Mark G. Volkovitsh Zoological Institute Russian Academy of Sciences Universitetskaya nab. 1 199034 St. Petersbourg, Russia e-mail: [email protected]

Bernhard J. v. Vondel Natuurmuseum Rotterdam Roestuin 78 3343 CV Hendrik-Ido-Ambacht The Netherlands e-mail: [email protected] Kipling Will ESPM-Division of Insect Biology University of California 201 Wellman Hall Berkeley, CA 94720-3112, USA e-mail: [email protected]

1. Systematic position, basal branching pattern and early evolution Rolf G. Beutel

The Coleoptera is one of 11 conventionally recognised orders of holometabolous insects (Kristensen 1999). Their position within Endopterygota is indicated by the presence of a pupal stage, absence of external wing and genital-appendage buds in the larval stages, and distinctive larval eyes, the stemmata. As in other groups of holometabolous insects, the larvae differ strongly from adults in terms of morphology and life-habits. A close relationship with Neuropterida was proposed by Hennig (1969), Mickoleit (1973), and others. Possible synapomorphies shared by beetles and the neuropteroid orders were summarised by Lawrence & Newton (1982) as follows (a) presence of a gula in the adult stage (secondarily absent in lacewings), (b) an oblique attachment of the forewings with an enlargement of the costal field (humeral and epipleural areas of the elytron), (c) transformation of the ovipositor (Mickoleit 1973), and (d) structure of the stemmata. A clade comprising Neuropterida and Coleoptera is also supported by the findings of Afzelius & Dallai (1994): the sperm axoneme of Coleoptera and the neuropteroid orders is characterised by intertubular material that is divided into two very specific portions. Additionally four synapomorphic features of the meta wing base were proposed by Hörnschemeyer (1998). Close affinities between Neuropterida and Coleoptera were confirmed in a recent work on attachment structures of insect legs and the ordinal phylogeny of Hexapoda (Beutel & Gorb 2001), and can be considered as reasonably well established. However, in all minimal length trees presented in this study, Strepsiptera was placed as closest relative of Coleoptera. The clade Neuropterida + (Coleoptera + Strepsiptera) is supported by three unambiguous apomorphies: (a) enlarged head of axillary I (54.1; Hörnschemeyer 1998), (b) caudal process of axillary II (55.1; Hörnschemeyer 1998), and (c) gonocoxite fused with stylus base (75.1; ovipositor secondarily reduced in strepsipteran females). The presence of hairy ventral surfaces of the tarsomeres is another potential synapomorphy (109.1). However, this interpretation implies secondary loss in neuropterans, and specific attachment hairs are almost certainly absent from the groundplan of Strepsiptera (Mengenillidae; Pohl & Beutel 2004). Potential synapomorphies of Strepsiptera + Coleoptera are the loss of M. (= muscle) furco-pleurocostales III (58.1) and M. scutello-postnotalis III (59.1), and

the absence of dorsal pulsatile organs. Posteromotorism and the decrease in size of the mesothorax were excluded from the data matrix as unspecific character states, as was the loss of mesothoracic muscles, which is obviously correlated with posteromotorism. A close relationship between Coleoptera and Strepsiptera contrasts with a hypotheses presented by Whiting et al. (1997) and Wheeler et al. (2001). However, the results presented by these authors are not fully consistent. Their cladistic analysis of morphological data resulted in a sister-group relationship between Strepsiptera and Antliophora, and a clade Strepsiptera + Diptera (= "Halteria") was supported when all data (including 18S- and 28S rDNA sequences) were analysed. It has to be pointed out, that the morphological data matrices presented by Whiting et al. (1997) and Wheeler et al. (2001) contain some problematic codings (s. Beutel & Gorb 2001), and the proposed antliophoran-strepsipteran synapomorphies are not convincing due to homplasy (e. g., reduction of labial palp segments) or are absent in the groundplan of Strepsiptera (dagger-like mandibles; H. Pohl, pers. comm.). In the molecular analyses (Wheeler 2001), many generally recognised groups of insects were rendered non-monophyletic. Strepsiptera are not close to Diptera in the 28S r D N A tree, and are not the sistergroup of Diptera in the 18S rDNA tree. In the latter cladogram, Strepsiptera + Metajapyx (Diplura) is the sistergroup of an assemblage comprising the dipteran genera and Myrmeleon immaculatus (Neuroptera). It is obvious, that a reliable positioning of Strepsiptera needs further study (Kristensen 1999), and a close relationship of Coleoptera with this group is far from well established. Several plesiomorphic conditions suggest an isolated position of Strepsiptera, possibly as sistergroup of all remaining endopterygote orders. Larval external wing buds (Mengenillidae) and compound eyes occur, and their development differs strongly from other holometabolous insects (H. Pohl, pers. comm.). Strepsipterans lack a typical pupa and are characterised by hypermetamorphosis. The changes between immature stages are more dramatic than those taking place in the final transformations (H. Pohl, pers. comm.). The presence of a well developed abdominal segment XI in first instar larvae (Pohl 2000) is another feature distinguishing Strepsiptera from all other groups of Endopterygota.

Rolf G. Beutel Archostem.s.str. Adephaga

Tertiary

Myxophaga

Polyphaga

= Schizophor. ι

\ i JJ = Ademosyn.

= Catiniidae

Cretaceous

Jurassic

= Triadocup.

I TI rriassic iaooio

= Proto-= Permo- =Rhomco|

cup

32.1

9

29.1-31.1

I

Γ

boc

»19.1-24.1 17.1 16.1 1

I 11.1

Permian

14.1-15.1

112.1-13.1 1.1-10.1

Carboniferous Fig. 1.1. Ciadogram showing interrelationships of fossil and extant groups of Pan-Coleoptera (Beutel 1997). Apomorphies: 1.1 — Elytra, 2.1 - metathorax enlarged, sternum longer than tergum, 3.1 - alae at least partly covered by elytra at rest, 4.1 —no exposed membranes, 5.1 — broad prothoracic postcoxal bridge, 6.1 — metacoxae transverse and flat, movability restricted, 7.1 — abdominal sternite I reduced, 8.1 — coxosternite VIII retracted, 9.1 — tuberculate surface structure of cuticle, 10.1 —13 antennomeres, 11.1 — ovipositor unpaired, acuminate, 12.1 —elytra not broader than abdominal segments, 13.1 - elytra with regular arrangement of unsclerotized cells, 14.1 - 11 antennomeres, 15.1 - elytra as long and as broad as abdomen, subelytral chamber, 16.1 — procoxal cavity open, 17.1 prosternal process narrow, apically rounded, 18.1 — tergite IX of larvae with urogomphi.

The monophyly of Pan-Coleoptera ( = Coleoptera sensu lato·, s. Lauterbach 1989 1 ) is supported by several autapomorphies (Beutel 1997): (a) forewings transformed into sclerotized elytra with epipleura; (b) cuticle strongly sclerotized and exposed membranes absent; (c) head prog1

A Pan-monophylum as defined by Lauterbach (1989) includes all extant species and all stem-lineage representatives, i. e. all fossil taxa which split off before the first extant subgroup.

nathous, flattened and wedge-shaped; (d) antennae shortened, 13-segmented (Protocoleoptera and Permocupedidae; Ponomarenko 1969); (e) meta wings folded longitudinally in resting position; (f) flight function largely or completely restricted to metathorax; (g) metathorax enlarged, ventrite longer than tergite, (h) abdominal segment I completely reduced; (i) prothorax with prosternal process and broad postcoxal bridge (Protocoleoptera, Permocupedidae, Rhombocoleidae; Ponomarenko 1969); (j) mesothoracic

Systematic position, basal branching pattern and early evolution meron absent; (k) metacoxae broadened and flattened, moveability largely restricted to adduction and abduction; (1) posterior abdominal segments invaginated, sternite VII terminal; and m) cuticle with tubercular surface structure. Protocoleoptera ( = Tshekardocoleidae; Crowson 1975; Kukalovä-Peck 1991) are characterised as a monophyletic group by a pointed, unpaired structure at the abdominal apex (Fig. 1.1: 11.1), which may represent an unusual type of ovipositor (see Kukalovä 1969: Fig. 1; Ponomarenko 1969: Fig. 29 b). A sister-group relationship with the remaining Pan-Coleoptera was proposed by Beutel (1997). The monophyly of Pan-Coleoptera excluding Protocoleoptera is supported by modifications of the sclerotized forewings. The elytra are better adapted to the shape of the abdomen as is the case for the tshecardocoleids (Fig. 1.1: 12.1; s. Kukalovä 1969: Figs. 1, 5; Kukalovä-Peck 1991). The original venation of the fore-wings is still partly preserved in Protocoleoptera (Kukalovä-Peck 1991: Fig. 6.28 B), whereas adults of all other groups of Coleoptera are characterised by a regular arrangement of the unsclerotized cells or by smooth elytra (Fig. 1.1: 13.1). Taldycupedidae (L. Permian-U. Jurassic; Ross & Jarzembowski 1993), Asiocoleidae (Permian; Ross & Jarzembowski 1993) and Tricoleidae (Triassic - Jurassic; Ross & Jarzembowski 1993) belong to Pan-Coleoptera. This is documented by the presence of elytra with epipleura. However, a more precise systematic placement would be hazardous at present, as other body parts are largely unknown. Pan-Coleoptera, excluding Protocoleoptera and Permocupedidae, is a monophyletic group characterised by a subelytral chamber, which results from the complete morphological adaptation of the elytra to the abdomen (Fig. 1.1: 15.1). Another apomorphy of this clade is the loss of 2 antennal segments (Fig. 1.1: 14.1). A flagellum with 9 segments is present in Rhombocoleidae and adults of most extant groups of Coleoptera. Pan-Coleoptera excluding Protocoleoptera, Permocupedidae and Rhombocoleidae are characterised by the loss of the prothoracic postcoxal bridge (Fig. 1.1: 16.1). Postcoxal bridges appear secondarily within some taxa of several groups of extant Coleoptera such as Carabidae and Hydraenidae. The monophyly of Coleoptera (sensu stricto) is largely undisputed, even though it is not supported by all molecular data (e.g., Caterino et al. 2002). The monophyly of a clade comprising Colpocaccus (Adephaga, Carabidae), Priacma serrata (Archostemata, Cupedidae), Megaloptera and Raphidioptera ( " M R P C clade") (Whiting et al. 1997) was an artefact resulting from contamination (Wheeler et al. 2001). A possible autapo-

3

morphy of extant beetles (Pan-Coleoptera excl. Protocoleoptera, Permocupedidae, Rhombocoleidae, and Triadocupedinae) is the narrowed prosternal process (Beutel 1997; Fig. 1.1: 17.1). Monophyly was also clearly supported by 23 autapomorphic features in a cladistic analysis of a broad characterset of extant representatives (Fig. 1.2; Beutel & Haas 2000) namely: sclerites firmly connected (20.1); eleven antennomeres (24.1); pre-episternal cervical sclerites absent (31.0); pronotum with inflected margin (32.1); M. cervicale-occipitalis torquatus absent (40.1); M. pleura-trochanteralis present (44.0); mesothoracic meron absent (50.1); scutellar elytralocking device present (52.1); M. scutello-postnotalis absent (53.1); M. intraepisternalis II absent (57.1); M. pleurocosto-praenotalis absent (61.1); metascutal fissure present (67.1); alacristae present (68.1); metacoxae transverse (71.1), less than five costal cross-veins (73.0), meta wings folded under elytra (74.1); apical part of wings rolled (75.1); triangular fold of alae present, proximal to R A 3 + 4 (81.1); M. noto-pleuralis b III absent (91.1); Mm. furco-pleurocostales III absent (93.1); M. sterno-coxalis III absent (96.1); sternite I strongly reduced (100.1); and four malpighian tubules (104.2). The basal branching pattern within Coleoptera is controversial. Archostemata were considered as the sister-group of the remaining Coleoptera, and Adephaga as the sister-group of a clade comprising Polyphaga and Myxophaga by Crowson (1955). Sister-group relationships between (Adephaga + Archostemata) and (Polyphaga + Myxophaga), respectively, were proposed by Baehr (1979), based on a comparative study of prothoracic structures. A branching pattern of Polyphaga + (Archostemata + (Adephaga + Myxophaga)) was suggested by Kukalovä-Peck & Lawrence (1993) and KukaloväPeck & Lawrence (2004). The following character states were considered as presumptive apomorphies of Coleoptera excluding Polyphaga (Kukalovä-Peck & Lawrence 1993): (a) central field of alae triangular and forming a pocket; (b) medial bar with distinct, abrupt hinge; (c) basal portion of R A 3 + 4 cut twice by the triangular fold; (d) CuP~ completely lost; (e) anterior anal basivenale (BAA) present and V-shaped; (f) characteristic 1 Ax with a massive, broad head and very long notched tail; and (g) cubital proxalare (PRCu) fused to the tergum. Four synapomorphies were proposed for Adephaga + Myxophaga: (a) R A 3 + 4 delimiting the radial cell posteriorly only to about the middle of the cell, where the vein divides; (b) cross vein r3 absent and r4 shifted proximally; (c) R P forming a wide angle with RA, so that radial and central fields are very large; and (d) A P 3 + 4 unbranched. The polarity determination in this study was not based on a cladistic analysis of the data.

4

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