THE BIOLOGY OF TERRESTRIAL MOLLUSCS This Page Intentionally Left Blank THE BIOLOGY OF TERRESTRIAL MOLLUSCS Edited by G.M. Barker Landcare Research Hamilton New Zealand CABI Publishing CABI Publishing is a division of CAB International CABI Publishing CABI Publishing CAB International 10 E 40th Street Wallingford Suite 3203 Oxon OX10 8DE New York, NY 10016 UK USA Tel: +44 (0)1491 832111 Tel: +1 212 481 7018 Fax: +44 (0)1491 833508 Fax: +1 212 686 7993 Email: [email protected] Email: [email protected] © CAB International 2001. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data The biology of terrestrial molluscs/edited by G.M. Barker. p. cm. Includes bibliographical references. ISBN 0-85199-318-4 (alk. paper) 1. Mollusks. I. Barker, G.M. QL407 .B56 2001 594--dc21 00-065708 ISBN 0 85199 318 4 Typeset by AMA DataSet Ltd, UK. Printed and bound in the UK by Cromwell Press, Trowbridge. Contents Contents Contents Contributors vii Preface ix Acronyms xi 1 Gastropods on Land: Phylogeny, Diversity and Adaptive Morphology 1 G.M. Barker 2 Body Wall: Form and Function 147 D.L. Luchtel and I. Deyrup-Olsen 3 Sensory Organs and the Nervous System 179 R. Chase 4 Radular Structure and Function 213 U. Mackenstedt and K. Märkel 5 Structure and Function of the Digestive System in Stylommatophora 237 V.K. Dimitriadis 6 Food and Feeding Behaviour 259 B. Speiser 7 Haemolymph: Blood Cell Morphology and Function 289 E. Furuta and K. Yamaguchi v vi Contents 8 Structure and Functioning of the Reproductive System 307 B.J. Gómez 9 Regulation of Growth and Reproduction 331 A. Gomot de Vaufleury 10 Spermatogenesis and Oogenesis 357 J.M. Healy 11 Population and Conservation Genetics 383 T. Backeljau, A. Baur and B. Baur 12 Life History Strategies 413 J. Heller 13 Behavioural Ecology: On Doing the Right Thing, in the Right Place at the Right Time 447 A. Cook 14 Soil Biology and Ecotoxicology 489 R. Dallinger, B. Berger, R. Triebskorn-Köhler and H. Köhler Index 527 Contributors Contributors Contributors T. Backeljau, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, B-1000 Brussels, Belgium G.M. Barker, Landcare Research, Private Bag 3127, Hamilton, New Zealand A. Baur, Department of Integrative Biology, Section of Conservation Biology (NLU), University of Basel, St Johanns-Vorstadt 10, CH-4056 Basel, Switzerland B. Baur, Department of Integrative Biology, Section of Conservation Biology (NLU), University of Basel, St Johanns-Vorstadt 10, CH-4056 Basel, Switzerland B. Berger, Institut für Zoologie und Limnologie, Abteilung Ökophysiologie, Universität Innsbruck, Technikerstraße 25, A-6020 Innsbruck, Austria R. Chase, Department of Biology, McGill University, 1205 Av. Docteur Penfield, Montréal, Québec, Canada H3A 1B1 A. Cook, School of Environmental Studies, University of Ulster, Coleraine, Northern Ireland BT52 1SA, UK R. Dallinger, Institut für Zoologie und Limnologie, Abteilung Ökophysiologie, Universität Innsbruck, Technikerstraße 25, A-6020 Innsbruck, Austria I. Deyrup-Olsen, Department of Zoology, University of Washington, Seattle, Washington, USA V.K. Dimitriadis, School of Biology, Faculty of Sciences, Aristotle University of Thessaloniki, Thessaloniki, Greece 54006 E. Furuta, Department of Histology, Dokkyo University School of Medicine, Mibu, Tochigi 321-0293, Japan vii viii Contributors B.J. Gómez, Departamento de Zoología y Dinámica Celular Animal, Facultad de Farmacia, Universidad del País Vasco, Paseo de la Universidad 7, 01006 Vitoria, Spain A. Gomot de Vaufleury, Laboratoire de Biologie et Ecophysiologie, Faculté des Sciences et des Techniques, Université de Franche-Comté, Place Leclerc, 25030 Besançon Cedex, France J.M. Healy, Centre for Marine Studies, The University of Queensland, Brisbane, Queensland 4072, Australia J. Heller, Department of Evolution, Systematics and Ecology, The Hebrew University, Jerusalem 91904, Israel H. Köhler, Animal Physiological Ecology, Zoological Institute, Konrad-Adenauer-Strasse 20, D-72072 Tübingen, Germany D.L. Luchtel, Department of Environmental Health, School of Public Health and Community Medicine, University of Washington, Box 357234, Seattle, WA 98195-7234, USA U. Mackenstedt, Institut für Zoologie, Fachgebiet Parasitologie, Universität Hohenheim, Emil-Wolff-Str. 34, D-70599 Stuttgart, Germany K. Märkel, Lehrstuhl für Spezielle Zoologie, Ruhr-Universität Bochum, Universitätstraße 150, D-44780 Bochum, Germany B. Speiser, Research Institute of Organic Agriculture (FiBL), Ackerstrasse, CH-5070 Frick, Switzerland R. Triebskorn-Köhler, Steinbeis-Transfer Center Ecotoxicology and Ecophysiology, Kreuzlinger Strasse 1, D-72108 Rottenburg, Germany and Animal Physiological Ecology, Zoological Institute, Konrad-Adenauer-Strasse 20, D-72072 Tübingen, Germany K. Yamaguchi, Institute of Medical Science, Dokkyo University School of Medicine, Mibu, Tochigi 321-0293, Japan Preface Preface Preface With an estimated 35,000 species, terrestrial gastropod molluscs are one of the most successful and diverse animal groups in land-based eco- systems. These animals have long been of importance to human societies as food, medicine, crop pests, vectors of parasites, and as tools, personal ornamentation and currency in trade. A relatively small fraction of the global terrestrial gastropod diversity – in the order of 100 species – has proved to be highly adaptive to environmental change brought about by human activity, and often become a highly abundant and characteristic component of invertebrate faunas in modified habitats. These species generally exhibit high propensities for both passive dispersal associated with human trading activities and invasiveness when introduced to new areas. They have become increasingly important as crop pests in agriculture and as vectors of helminth parasites in humans and domestic livestock. Because of their increasing ubiquity and economic importance, and their utility as laboratory animals, these same species are among the most intensively studied invertebrates. They have proved to be excellent model systems for studies in, for instance, neurophysiology, behavioural ecology and population genetics. By virtue of their capacity to accumulate metal in their tissues, these animals are also of increasing utility as bioindicators of environmental pollution. Many of these areas of investi- gation show that research on terrestrial gastropods can contribute to the wider fields of research endeavour encompassing physiology, ecology, evolution and conservation biology. The greater part of the terrestrial gastropod biodiversity, however, goes unnoticed by people – they are mostly small creatures, seeking out a living as detritivores, contributing significantly to nutrient cycling through facilitation of decomposition and return of plant litter to the soil. In many parts of the world, terrestrial gastropod communities are being ix x Preface greatly perturbed or lost through human-induced habitat degradation and loss. This book presents a synthesis of current knowledge and research on the biology of terrestrial gastropod molluscs. In a series of peer-reviewed chapters, it provides authoritative coverage of the topics of morphology, phylogeny and systematics, structure and function of the various organ systems, feeding behaviour, life history strategies, behavioural ecology, population and conservation genetics, and soil biology and ecotoxicology. This book is for both students and professionals concerned with terrestrial Mollusca. Gary M. Barker Landcare Research Hamilton New Zealand Acronyms Acronyms Acronyms ABARE Australian Bureau of Agricultural and Resource Economics ACIAR Australian Council for International Agricultural Research ACSAD Arab Centre for Studies of Arid Zones and Dry Lands ANZECC Australian and New Zealand Environment and Conservation Council AOAD Arab Organization for Agricultural Development ATO African Timber Organization BD Bilateral donors CAI Current annual increment CATIE Tropical Agricultural Research and Higher Education Centre CBD Convention on Biological Diversity CCAB-AP Central American Council on Forests and Protected Areas CCAD Central American Commission on Environment and Development CCC Convention on Climate Change CCD Convention to Combat Desertification CCFM Canadian Council of Forest Ministers CFTT Centre for Forest Tree Technology C&I Criteria and indicators CIFOR Center for International Forestry Research CPOM Coarse particulate organic matter CRPF Centre Régional de la Propriété Forestière CSA Canadian Standards Association CSCE Conference on Security and Cooperation CSD Commission on Sustainable Development CWD Coarse woody debris DEST Department of Environment, Sport and Tourism (Aust.) DFID Department for International Development (UK) xi xii Acronyms DNRE Department of Natural Resources and Environment (Vic.) DPCSD Department for Policy Coordination and Sustainable Devel- opment (UN) ECE Economic Commission for Europe EFI European Forestry Institute EMS Environmental management system ENGO Environmental non-government organization ESFM Ecologically sustainable forest management EU European Union Fa Facilitator FAO Food and Agriculture Organization FLORES Forest Land Oriented Resource Envisioning System FMU Forest management unit FRA 2000 Forest Resources