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Arthropod Relationships

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The Systematics Association Special Volume Series

Series Editor

David M. John

Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD, UK

The Systematics Association provides a forum for discussing systematic problems and integrating new information from cytogenics, ecology and other specific fields into taxonomic concepts and activities. It has achieved great success since the Association was founded in 1937 by promoting major meetings covering all areas of biology and palaeontology, supporting systematic research and training courses through the award of grants, production of a membership newsletter and publication of review volumes by its publisher Chapman & Hall. Its membership is open to both amateurs and professional scientists in all branches of biology who are entitled to purchase its volumes at a discounted price.

The first ofthe Systematics Association's publications, The New Systematics, edited by its then president Sir Julian Huxley, was a classic work. Over 50 volumes have now been published in the Association's 'Special Volume' series often in rapidly expanding areas of science where a modem synthesis is required. Its modus operandi is to encourage leading exponents to organise a symposium with a view to publishing a multi-authored volume in its series based upon the meeting. The Association also publishes volumes that are not linked to meetings in its 'Volume' series.

Anyone wishing to know more about the Systematics Association and its volume series are invited to contact the series editor. Further information about the Systematics Association can also be found on the website

http://www.thomson.com/systematic/SAImain.html

Forthcoming titles in the series:

Evolutionary Relationships Among Protozoa Coombs et al.

Advances in Plant Molecular Systematics Bateman et al.

Homology in Systematics R.W. Scotland and R.T. Pennington

The Flagellates B. Leadbeater and J. Green

Other Systematics Association publications are listed after the index for this volume.

The Systematics Association Special Volume Series 55

Arthropod Relationships

Edited by

R.A. Fortey Department of Palaeontology, The Natural History Museum, London, UK

and

R.R. Thomas Department of Zoology, The Natural History Museum, London, UK

I ~I' I SPRINGER-SCIENCE+BUSINESS MEDIA, B. V.

First edition 1998

© 1998 Springer Science+Business Media Dordrecht

Origina11y published by The Systematics Association in 1998 Softcover reprint ofthe hardcover Ist edition 1998

Typeset in 10/12pt Times by Saxon Graphics Ltd, Derby

ISBN 978-94-010-6057-8 ISBN 978-94-011-4904-4 (eBook) DOI 10.1007/978-94-011-4904-4

Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the UK Copyright Designs and Patents Act, 1988, this publication may not be reproduced, stored, or transmitted, in any form or by any means, without the prior permis sion in writing of the publishers, or in the case of reprographic reproduction only in accordance with the terms of the licences issued by the Copyright Licensing Agency in the UK, or in accordance with the terms of licences issued by the appropriate Reproduction Rights Organization outside the UK. Enquiries conceming reproduction outside the terms stated here should be sent to the publishers at the London address printed on this page.

The publisher makes no representation, express or implied, with regard to the accuracy ofthe information contained in this book and cannot accept any legal responsibility or liability for any errors or omissions that may be made.

A catalogue record for this book is available from the British Library

Library of Congress Catalog Card Number: 97---68214

@ Printed on acid-free text paper, manufactured in accordance with ANSIINISO Z39.48-1992 (Permanence of Paper).

To the memory of Sidnie Manton, 20 years on from the publication of The Arthropoda.

CONTENTS

List of contributors

Preface

1 Bodyplans, phyla and arthropods J. W. Valentine and H Hamilton

2 The phylogenetic position of the Arthropoda C. Nielsen

3 A defence of arthropod polyphyly G. Fryer

4 Hox genes and annelid-arthropod relationships MHDick

5 Arthropod and annelid relationships re-examined D.J. Eernisse

6 Evolutionary correlates of arthropod tagmosis: scrambled legs MA. Wills, D.E.G. Briggs and R.A. Fortey

7 Theories, patterns, and reality: game plan for arthropod phylogeny MJ. Emerson and F.R. Schram

8 Sampling, groundplans, total evidence and the systematics of arthropods W. C. Wheeler

9 Arthropod phylogeny: taxonomic congruence, total evidence and conditional combination approaches to morphological and molecular data sets J. Zrzary, V. Hypsa and M Vlaskowl

10 The place of tardigrades in arthropod evolution R.A. Dewel and W. C. Dewel

11 Stem group arthropods from the Lower Cambrian Sirius Passet fauna of North Greenland G.E. Budd

12 Cambrian 'Orsten' -type arthropods and the phylogeny of Crustacea D. Walossek and K.J. Muller

13 Comparative limb morphology in major crustacean groups: the coxa-basis

joint in postmandibular limbs G. Boxshall

ix

Xl

11

23

35

43

57

67

87

97

109

125

139

155

Vlll

14 Crustacean phylogeny inferred from 18S rDNA T. Spears and L. G. Abele

Contents

15 A phylogeny of recent and fossil Crustacea derived from morphological characters MA. Wills

16 The fossil record and evolution of the Myriapoda WA. Shear

17 The early history and phylogeny of the chelicerates J.A. Dunlop and P.A. Selden

18 Problem of the basal dichotomy of the winged insects A.P. Rasnitsyn

19 Arthropod phylogeny and 'basal' morphological structures J. KukalowJ-Peck

20 Advances and problems in insect phylogeny R. Willmann

21 The groundplan and basal diversification of the hexapods N.P. Kristensen

22 Phylogenetic relationships between higher taxa of tracheate arthropods 0. Kraus

23 Myriapod-insect relationships as opposed to an insect-crustacean sister group relationship W Dohle

24 Cleavage, germ band formation and head segmentation: the ground pattern of the Euarthropoda G. Scholtz

25 Homology and parallelism in arthropod sensory processing D.-E. Nilsson and D. Osorio

26 The organization and development of the arthropod ventral nerve cord: insights into arthropod relationships P.M Whitington and J.P. Bacon

Index

169

189

211

221

237

249

269

281

295

305

317

333

349

371

CONTRIBUTORS

L.G. ABELE Office of the Provost, Florida State University,

Tallahassee, FL 32306-3020, USA email: [email protected]

J.P. BACON Sussex Centre for Neuroscience, School of Biological

Sciences, University of Sussex, Brighton BN1 9QG, UK email: [email protected]

G. BOXSHALL Department of Zoology, The Natural History Museum,

Cromwell Road, London, SW7 5BD, UK email: [email protected]

D.E.G. BRIGGS Department of Geology, Wills Memorial Building,

University of Bristol, Queen's Road, Bristol, BS8 1RJ, UK

email: [email protected]

G.BUDD Department of Historical Geology and Palaeontology,

Institute of Earth Sciences, Norbyvagen 22, S-75236 Uppsala, Sweden


R.A. DEWEL Department of Biology, Appalachian State University,

Boone, NC 28608, USA email: [email protected]

W.C.DEWEL Department of Biology, Appalachian State University,

Boone, NC 28608, USA email: [email protected]

M.H.DICK Department of Biology, Middlebury College, Middlebury,

Vermont 05753, USA email: [email protected]

W.DOHLE Institut fiir Zoologie, Freie Universitat Berlin, Konigin

Luise-Str. 1-3, D-14l95 Berlin, Germany

J.A. DUNLOP Department of Earth Sciences, University of Manchester,

Oxford Road, Manchester,M13 9PL, UK email: [email protected]

D.J. EERNISSE Department of Biology MH282, California State

University, Fullerton, CA 92834, USA email: [email protected]

M.J. EMERSON (Deceased) late of the San Diego Natural History Museum

R.A. FORTEY Department of Palaeontology, The Natural History

Museum, Cromwell Road, London, SW7 5BD, UK email: [email protected]

G.FRYER Institute of Environmental and Biological Sciences,

University of Lancaster, Lancaster LA1 4QY, UK

H. HAMILTON Department of Integrative Biology, Museum of

Paleontology, University of California, Berkeley, CA 94720, USA

V.HYPSA Faculty of Biological Sciences, Branisovska 31, 370 05

Ceske Budejivce, Czech Republic

O.KRAUS Zoologisches Institut und Zoologisches Museum,

Universitiit Hamburg, Martin-Luther-King-Platz 3, D-20146 Hamburg, Germany

N.P. KRISTENSEN Zoological Museum, University of Copenhagen,

Universitetsparken 15, DK-2l00 Copenhagen 0, Denmark


J. KUKALOV A-PECK Department of Earth Sciences, Carleton University, 1125

Colonel By Drive, Ottawa, KI S 5B6 Canada


x Contributors

K.J. MULLER Institiit fUr PaHiontologie, Universitiit Bonn, Nussallee 8,

D-53115 Bonn, Germany

C. NIELSEN Zoological Museum, University of Copenhagen,

Universitetsparken 15, DK-2100 Copenhagen 0, Denmark


D.-E. NILSSON Department of Zoology, University of Lund,

Helgonavagen 3, S-223 62 Lund, Sweden email: [email protected]

D. OSORIO Biological Sciences, Sussex University, Brighton,

BNI9QG, UK email: [email protected]

AP. RASNITSYN Arthropoda Laboratory, Paleontological Institute, Russian

Academy of Science, Profsoyuznaya Str. 123, Moscow,117647, Russia


G. SCHOLTZ Institut fUr Biologie, Vergleichende Zoologie Humboldt

Universitiit zu Berlin, Philippstr. 13, D-10115 Berlin, Germany


F.R. SCHRAM Institute for Systematics and Population Biology,

University of Amsterdam, Post Box 94766, NL-IOO GT Amsterdam, The Netherlands


P.A SELDEN Department of Earth Sciences, University of Manchester,

Oxford Road, Manchester, M13 9PL, UK email: [email protected]

W.A. SHEAR Department of Biology, Hampden-Sydney College,

Hampden-Sydney, VA 23943, USA email: [email protected]

T. SPEARS Department of Biological Sciences, Florida State

University, Tallahassee, FL 32306-2043, USA email: [email protected]

R.H. THOMAS Department of Zoology, The Natural History Museum,

Cromwell Road, London, SW7 5BD, UK email: [email protected]

J.W. VALENTINE Department ofIntegrative Biology, Museum of

Paleontology, University of California, Berkeley, CA 94720, USA


M. VLAsKOvA Faculty of Biological Sciences, Branisovska 31,37005

Ceske Budejovce, Czech Republic email: [email protected]

D. WALOSSEK Sektion fUr Biosystematische Dokumentation, University

ofUlm, Liststral3e 3, D-89079 UIm, Germany email: [email protected]

W.C. WHEELER Department ofInvertebrates, American Museum of Natural

History, Central Park West at 79th Street, New York, NY 10024-5192, USA


P.M. WHITINGTON Department of Zoology, University of New England,

Armidale, NSW, 2351, Australia email: [email protected]

R. WILLMANN II. Zoologisches Institut der Universitat, Georg-August

Universitat-G6ttingen, Berliner Stral3e 28, D-37073 G6ttingen, Germany

M.A WILLS Department of Geology, Wills Memorial Building,

University of Bristol, Queen's Road, Bristol BS8 lRJ, UK

email: [email protected]. uk

J. ZRZAvY Faculty of Biological Sciences, Branisovska 31,37005

Ceske Budejovce, Czech Republic email: [email protected]

PREFACE

Arthropods - insects, crustaceans, myriapods and arachnids - are the most speciose of all animal groups, and have probably been so for hundreds of millions of years. Their importance in every ecosystem - terrestrial and marine - is not in question. Yet very little has been agreed about how they achieved their pre-eminence. The evolutionary pathways which led to their current diversity are still the subject of controversy, despite the fact that many of the questions of descent have been debated for more than a century. We judged that the time was ripe for a new attempt at a synthesis of current views about arthropod phylogeny, the first since Gupta (1979). There have been many new advances in both methodology and technology since then, not least the impact of cladistics in the former, and molecular biology in the latter. Yet the classical approaches of comparative morphology, embryology, and palaeontology have been far from static - for example, there have been more discoveries of exciting new fossils in the past 20 years than ever before. The way forward seemed to be to bring together the scattered specialists to discuss a modem overview on arthropod relationships. Hence, we invited the leading figures in the various disciplines ofarthropodology to a Symposium at the Natural History Museum, London, 17-19 April 1996. We were generously supported by the Linnean Society of London, the Systematics Association, The Royal Society and the Natural History Museum in making the meeting a practical proposition, and to these bodies we tender our thanks. This book is the outcome of three days of mutual education.

We have taken a novel approach to publication of data. Many of the papers include cladistic analyses based upon character matrices - both morphological and molecular. We have published the results of the analyses in the book, but the data are stored and available on a website page. They can therefore be directly downloaded by other investigators, and manipulated as they choose. We trust that this will further aid the utility of the book.

The papers are arranged in such a way as to focus progressively from general problems to issues involving particular groups. The place of the Arthropoda within the animal phyla is considered first, particularly the relationship to annelids; there are reviews of the morphological, developmental and molecular data bearing on the unsolved problems. Most contributors favour arthropod monophyly, although Fryer bids us look carefully at some of the supposed homologies. Next, cladistic treatments of the arthropods as a whole - from several theoretical standpoints - provide a summary of the issues dealt with in subsequent chapters. The reader will be able to judge whether combining all phylogenetic data sources in such analyses is more persuasive than trying to identity uniquely derived apomorphies as a basis for the definition of groups. The important place of the tardigrades within the arthropod stem-lineage is shown by Dewel and Dewel. That Cambrian fossils offer unique evidence in fleshing out the early history of arthropods is illustrated with reference to new Lower Cambrian animals from Greenland, and the celebrated 'Orsten' arthropods from Sweden and elsewhere. It is clear that some of the main lineages of arthropod radiation had already been distinguished by the early Cambrian. There is recent evidence that the antiquity of the main branches in the group may be even older (Fortey et aI., 1996; Wray et aI., 1996). No doubt this antiquity exacerbates the problems in unequivocally recognizing fundamental branching events.

Even within each major arthropod group there are problems in recognizing the relationships between major taxa. Three different attitudes to such problems in Crustacea are exemplified by the morphological, molecular and 'key character' approaches, respectively. They yield interesting differences. The fossil history of myriapods and chelicerates is reviewed. The reality or otherwise of the 'Uniramia' (myriapods plus insects) is discussed by several authors; recent reconsideration of morphological evidence fmds more support for an insect-{;rustacean relationship than has been claimed in recent years (also A verof and Cohen, 1997). This is one example where molecular phylogenies have led to a re-evaluation of classical evidence. Within the section on hexapods, the reader will find clear accounts of the several schools of thought regarding the most fundamental subdivisions of the greatest group of arthropods. The four papers on this topic provide an object lesson in the problems of interpretation of characters in unscrambling important, but ancient events. On the one hand it is undoubtedly true that, as KukalovaPeck claims, there are morphologies preserved in fossils which no longer survive, and which must surely bear on the phylogeny. On the other hand, the interpretation of the fossil characters has proved difficult, and there are cases where ambiguous morphology has been placed at the service of a favoured theory, with the attendant dangers of circularity.

Finally, embryological and physiological evidence is brought into the argument. There are examples in the sensory 'wiring' across arthropod groups of such striking similarity that it seems difficult to comprehend that they may have evolved in parallel.

xu Preface

If there is a general message to be drawn from this compilation, it is that all fields of endeavour have much to contribute to an understanding of arthropod relationships. It would be unwise to neglect the broad range of discoveries in favour of those from a narrow specialization alone. It will be clear that there is, as yet, no 'right' answer about arthropod phylogeny; but the journey towards it gets increasingly interesting.

REFERENCES

Averof, M. and Cohen, S.M. (1997) Evolutionary origin of insect wings from ancestral gills. Nature, 385, 627-30.

Fortey, R.A., Briggs, D.E.G. and Wills, M.A. (1996) The Cambrian evolutionary 'explosion': decoupling cladogenesis from morphological disparity. Biological Journal of the Linnean Society, 57, 13-33.

R.A.F. and R.H T.

Gupta, A.P. (1979) Arthropod Phylogeny, Van Nostrand Reinhold Co., New York.

Wray, G.A., Levinton, 1.S. and Shapiro, L.H. (1996) Molecular evidence for deep Precambrian divergences among metazoan phyla. Science, 274, 568-73.

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