Earthworm Ecology FROM DARWIN TO VERMICUL TURE FOR 1882. MAN ·I~ ·BVT ·A·woR...JV\· Frontispiece Cartoon from Punch, December 6th, 1881 Earthworm Ecology FROM DARWIN TO VERMICUL TURE Edited by J. E. Satchell Institute of Terrestrial Ecology Merlewood Research Station Grange-over-Sands Cumbria, UK LONDON NEW YORK CHAPMAN AND HALL First published 1983 by Chapman and Hall Ltd I I New Fetter Lane, London EC4P 4EE Published in the USA by Chapman and Hall 733 Third Avenue, New York NY100I7 © 1983 Chapman and Hall Ltd Softcover reprint of the hardcover 1st edition 2007 University Press, Cambridge ISBN-13: 978-94-009-5967-5 e-ISBN-13: 978-94-009-5965-1 DOl: 10.1007/978-94-009-5965-1 All rights reserved. No part ofthis book may be reprinted, or reproduced or utilized in any form or by any electronic, mechanical or other means, now known or hereafter invented, including photocopying and record­ ing, or in any information storage and retrieval system, without permission in writing from the Publisher. British Library Cataloguing in Publication Data Earthworm ecology. I. Opisthopora I. Satchell, J. E. 595.I' 46 QL39 I. 04 Library of Congress Cataloging in Publication Data Main entry under title: Earthworm ecology. Bibliography: p. Includes index. I. Opisthopora-Ecology. 2. Earthworm culture. I. Satchell, John E. QL39I.A6E2 7 1983 Contents Preface Xl Contributors xiii DARWIN'S CONTRIBUTION TO EARTHWORM ECOLOGY I Darwin's Formation of Vegetable Mould- its philo­ sophical basis M. S. Ghilarov I 2 D~rwin on earthworms - the contemporary back­ ground and what the critics thought O. Graff 5 EARTHWORMS AND ORGANIC MATTER 3 Darwin's 'vegetable mould' and some modern con­ cepts of humus structure and soil aggregation M. H. B. Hayes 19 4 Organic matter turnover by earthworms J. D. Stout 35 (with a section by B. J. O'Brien) 5 Effect of earthworms on the disappearance rate of cattle droppings P. Holter 49 6 Soil transport as a homeostatic mechanism for stabil­ izing the earthworm environment A. Kretzschmar 59 v Vl Contents EARTHWORM ECOLOGY IN GRASSLAND SOILS 7 Earthworm ecology in grassland soils J. K. Syers and J. A. Springett 67 8 Effect of earthworms on grassland on recently re­ claimed polder soils in the Netherlands M. Hoogerkamp, H. Rogaar and H. J. P. Eijsackers 85 (with Appendix by J. H. Oude Voshaar and H. J. P. Eijsackers) 9 The activities of earthworms and the fates of seeds J. D. Grant 107 EARTHWORM ECOLOGY IN CULTIVATED SOILS 10 Earthworm ecology in cultivated soils C. A. Edwards 123 I I Nitrogen turnover by earthworms in arable plots treated with farmyard manure and slurry N. C. Andersen 139 12 Earthworm population dynamics in different agricul­ tural rotations A. Lofs-Holmin 151 EARTHWORM ECOLOGY IN FOREST SOILS 13 Earthworm ecology in forest soil J. E. Satchell 161 14 Earthworm ecology in deciduous forests in central and southeast Europe A. Zicsi 171 EARTHWORM ECOLOGY IN TROPICAL AND ARID SOILS 15 Earthworms of tropical regions -some aspects of their ecology and relationships with soils K.E.L~ 179 16 The ecology of earthworms in southern Africa A. J. Reinecke 195 Contents Vll 17 Effects of fire on the nutrient content and microfiora of casts of Pheretima alexandri M. V. Reddy 209 EARTHWORMS AND LAND RECLAMATION 18 Earthworms and land reclamation J. P. Curry and D. C. F. Cotton 21 5 19 Earthworm ecology in reclaimed opencast coal mining sites in Ohio J. P. Vimmerstedt 229 20 Development of earthworm populations in aban­ doned arable fields under grazing management H. J. P. Eijsackers 241 EARTHWORMS AND SOIL POLLUTION 21 Heavy metal uptake and tissue distribution III earthworms M. P. Ireland 247 22 Heavy metals in earthworms in non-contaminated and contaminated agricultural soil from near Vancouver, Canada A. Carter, E. A. Kenney, T. F. Guthrie and H. Timmenga 23 Earthworms and TCDD (2,3,7,8-tetrachloro­ dibenzo-p-dioxin) in Seveso G. B. Martinucci, P. Crespi, P. Omodeo, G. Osella and G. Traldi 275 UTILIZATION OF WASTES BY EARTHWORM CULTURE 24 Earthworms as a source of food and drugs J. R. Sabine 285 25 Assimilation by the earthworm Eisenia jetida R. Hartenstein 297 viii Contents 26 The culture and use of Perionyx excavatus as a protein resource in the Philippines R. D. Guerrero 309 27 Utilization of Eudrilus eugeniae for disposal of cassava peel C. C. Mba 315 28 Cultivation of Eisenia jetida using dairy waste sludge cake K. Hatanaka, Y. Ishioka and E. Furuichi 323 29 The earthworm bait market in North America A. D. Tomlin 331 30 A simulation model of earthworm growth and popu- lation dynamics: application to organic waste con- verSIOn M. J. Mitchell 339 EARTHWORMS AND MICROFLORA 31 Earthworm microbiology J. E. Satchell 35 1 32 The effects of fungi on food selection by Lumbricus terrestris (L.) A. Cooke 365 33 Introduction of amoebae and Rhizobium japonicum into the gut of Eisenia jetida (Sav.) and Lumbricus terrestris L. J. Rouelle 375 34 Enterobacter aerogenes infection of Hoplochaetella suctoria B. R. Rao, 1. Karuna Sagar and J. V. Bhat 383 EARTHWORMS IN FOOD CHAINS 35 Predation on earthworms by terrestrial vertebrates D. W. Macdonald 393 Contents ix 36 Predation on earthworms by the Black-headed gull (Larus ridibundus L.) G. Cuendet 415 37 Agastrodrilus Omodeo and Vaillaud, a genus of car­ nivorous earthworms from the Ivory Coast P. Lavelle 425 EARTHWORM EVOLUTION AND DISTRIBUTION PATTERNS 38 The establishment of earthworm communities M. B. Bouche 431 39 The structure of earthworm communities P. Lavelle 449 TAXONOMY AND NOMENCLATURE 40 The scientific names of earthworms R. W. Sims 41 A guide to the valid names of Lumbricidae (Oligochaeta) E. G. Easton 475 Systematic index General index 492 Preface 'Darwin cleared: official' This 1982 Times (7 January) head­ line of a first leader, reporting the astonishing case brought in Arkansas against compulsory teaching of a biblical account of creation, hopefully set at rest doubts about Darwin in the minds of a public confused by media presentations of such unfamiliar concepts as punctuated equilibria, cladism and phenetics. Mud sticks, but Darwin's perturbed ghost may have found some consolation in the concurrent celebrations at Grange-over-Sands, a modest township in Cumbria, UK, of the centenary of the publication of his less controversial book The Formation of Vegetable Mould through the Action of Worms. In the form of a symposium on earthworm ecology, this attracted some 150 participants, predominantly adrenalin-charged research workers in the full heat of peer-group interaction. This book comprises a selection of the more ecologically oriented papers contributed to the symposium, brutally edited in the interests of brevity and thematic continuity. The book opens with an appraisal of Darwin's earthworm work in its historical and philosophical context and relates his views on 'vegetable mould' to current concepts of humus formation. Thereafter, quotations from Darwin made out of piety have been rigorously excluded. Subsequent sections each comprise a review chapter and two or three 'case studies' presenting new data on a related topic. It has always been difficult to formulate a balanced judgement on how far earthworm activity creates fertile soils and how far such soils create a favourable environment for earthworm activity. The early chapters of this book now facilitate a clear discrimination between the chicken and the egg and fully vindicate Xl XlI Preface Darwin's views on earthworm pedogenesis. Subsequent sections cover earthworm biology in natural and man-made ecosystems including polluted land and commercial vermiculture. No attempt has been made to impose uniformity in species nomenclature but the final chapters analyse this problem and propose a list of valid names as a basis for further attempts to resolve the current nomenclatural chaos. Each of the twelve review chapters cites some 20-100 published papers and together provide a rather comprehensive coverage of the earthworm ecology literature. They do not include references on earthworm physiology, enzyme chemistry, subjects like wound healing where the earthworm is of interest only as experimental material, or papers primarily of local distributional interest. A more comprehensive bibliography of the earthworm literature was prepared for distribution at the 1981 symposium and an updated supplement to this is now available. Stephenson's monograph 'The Oligochaeta' reviewed the earthworm literature up to 1930 and around 2500 papers have been published in scientific journals since. This does not include popular articles or the profuse manifestations of excessive drilophilia generated by commercial interests or organic faddists. Despite his well-known tag 'Natura non facit saltus', Darwin was quite clear that evolutionary rates are non-constant. Darwin's contemporaries thought of earthworms as pests to be eradicated. The publication of his book produced a great saltus in popular opinion, approaching modern concepts of the role of earthworms in soil formation, and with the subsequent doubling of the number of earthworm research papers every decade his ideas have now been fleshed out. Currently, the role of earthworms in many ecosystems is appreciated as being of much wider significance and of potential in such practical applications as pollution control and protein production. It is hoped that the publication of this book will provide a befitting tribute to a great pioneer of soil science and a base from which earthworm ecology can make a further leap forward. J. E. Satchell Contributors N. C. Andersen Zoology Department, Royal Veterinary and Agricultural University, Bulowsvej 13, DK-1870 Copenhagen V, Denmark. J. V. Bhat Laboratory of the Director of Research, Kasturba Medical College, Manipal, Pin-576119, India. M. B. Bouche Centre d'Etudes Phytosociologiques et Ecologiques, Laboratoire de Zooecologie du Sol, Route de Mende, BP 5051, F-34033 Montpellier Cedex, France.