Coleoptera, Beetles. Morphology and Systematics

Rolf G. Beutel, Richard A.B. Leschen, Rolf G. Beutel, Richard A. B. Leschen

• Publisher: De Gruyter
• Year: 2016
• ISBN:
  9783110386226
• Edition: 2
• Topic: Biological Sciences
• Subtopic: Entomology
• Index: Biological Sciences

This book is a revised edition of the first of three volumes in the Handbook of Zoology series which treats the systematics and biology of Coleoptera. With over 380, 000 described species, Coleoptera are by far the most species-rich order of insects and the largest group of animals of comparable geological age. Moreover, numerous species are tremendously important economically. The beetle volumes meet the demand of modern biologists seeking to answer questions about Coleoptera phylogeny, evolution, and ecology. This first Coleoptera volume covers the suborders Archostemata, Myxophaga and Adephaga, and the basal series of Polyphaga, with information on world distribution, biology, morphology of all life stages, phylogeny and comments on taxonomy.

Editors’ preface to the 2nd edition of Volume 1

Since the first of three Coleoptera volumes of the Handbook of Zoology series was published in 2005, the progress in the investigation of the most species-rich insect order has been remarkable. The exploration of the biodiversity of this extremely successful group of organisms has been dynamic, with valuable contributions made by individual naturalists, but also by research teams conducting large-scale biodiversity expeditions in different parts of the world. These efforts have yielded a remarkable increase in species numbers, as for instance in Hydrophiloidea from ca. 2,800 in 2005 (1st edition) to 3,335 in 2015 (1–12) (Volume 1 – chapter 12), or in Staphylinidae from 47,744 in 2005 to 55,224 in 2015 (1–14.7), the latter of which is also related to advances in classification. Moreover, new genera were described as well as two new families of Adephaga, which were not included in the 1st edition of Volume 1 (included in Volume 2 of 2010). The biodiversity explorations were accompanied by intensive taxonomic work conducted not only by professional researchers but also highly competent amateurs. Even though taxonomic research may not be greatly appreciated in parts of the scientific community, or governments, it is essential for the understanding of a highly complex and diverse group like Coleoptera, and generally for the development of stable classifications. An updated classification for Coleoptera is presented in Chapter 1–2. Important changes adopted here are the placement of Scirtoidea at the base of Polyphaga, and a new concept of Hydrophiloidea with subfamilies ranked as families (Chapter 1–12).

With remarkable technical innovations in insect anatomy in the last decade, the study of the morphology of Coleoptera has also intensified. Recently, as part of the Beetle Tree of Life project (funded by the United States National Science Foundation), an impressive morphological contribution was made by John Lawrence, Adam Ślipiński (CSIRO), and coworkers. This team of experts compiled, documented, and analyzed a comprehensive set of 516 characters representing 314 families and subfamilies of Coleoptera plus outgroups. New and detailed morphological data are available for species of many groups (e.g. cephalic and thoracic morphology of Archostemata, head morphology of Lepiceridae, Dascillidae, and Glaresidae, genital morphology of Ommatidae, e.g. 1–5.4), and larvae of several groups were described for the 1st time (Lepiceridae, 1–6.1, Meruidae, 1–7.8) or treated in detail morphologically, including complete 3D reconstructions of external and internal structures (e.g. 1st instar larvae of Tenomerga Neboiss [Cupedidae], 1–5.2).

The remarkable development of molecular systematics in the last decade had also a strong impact on beetle phylogenetics and evolutionary biology. A completely new chapter on molecular systematics is presented in this volume (1–3). One major achievement was the unambiguous systematic placement of Coleoptera as sistergroup to the highly specialized ectoparasitic Strepsiptera, both combined as Coleopterida, which in turn form the sistergroup to a clade containing the three neuropteroid orders. Another breakthrough was the identification of the scirtoid families and Derodontidae as basal lineages of the megadiverse Polyphaga.

The composition of the authors for this revised Volume 1 has changed slightly. Chuck Bellamy, regrettably passed away prior to the formulation of this new edition, and Buprestidae (1–18) were revised with the assistance of Mark Volkovitsch. Some authors were unavailable or were substituted by authors more recently active in the taxonomy of particular families. Duane McKenna is co-author of Systematic position and early evolution (1–1) and new author of Molecular systematics of Coleoptera (1–3), Lars Hendrich as new co-author of Dytiscidae (1–7.6), Pawel Jałoszyński as new author of Scydmaeninae (1–14.5), and Crystal Maier as leading author of Byrrhidae (1–19.1).

Not only are great strides being made to document beetle diversity and morphology, the investigation of beetle phylogeny is also presently progressing with great momentum. The phylogeny of the entire Coleoptera was addressed in the BTOL project and is presently investigated based on transcriptomes in a subproject of the 1KITE consortium (www.1KITE.org/, funded by Beijing Genomics Institute). This and other ongoing molecular investigations combined with new morphological findings and an intensified exploration of rich fossil material, will likely lead to a deeper understanding of beetle evolution and new classifications in the near future. The updated information compiled in this volume may facilitate and inspire ongoing research efforts, and new phylogenetic and evolutionary insights may eventually lead to new contributions in the Handbook of Zoology series.

Finally we would like to thank Kathleen Pruefer and Julia Reindlmeier for their support throughout the project.

Rolf G. Beutel and Richard A. B. Leschen

Editors’ preface to the 1st edition of Volume 1

With approximately 350 000 described species, Coleoptera are presently by far the most species rich 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 commonplace today. As 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 Adephaga, 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 including 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 80th 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 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 Volumes 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 recognize 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, 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 honored 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; FMNH; 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 support of this project.

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 and Richard A. B. Leschen

Contents

Editors’ preface to the 2nd edition of Volume 1

Editors’ preface to the 1st edition of Volume 1

Contributors

Rolf G. Beutel & Duane D. McKenna

1Systematic position, basal branching pattern and...