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Biology and Ecology of the Brown and Sea Trout Springer London Berlin Heidelberg New York Barcelona Hong Kong Milan Paris Santa Clara Singapore Tokyo J. L. Bagliniere, G. Maisse Biology and Ecology of the Brown and Sea Trout t Published in association with p Springer Praxis Publishing Chichester, UK Editeurs/Editors: J. 1. Bagliniere, G. Maisse INRA-ENSA, Station de Physiologie et Eeologie des Poissons 65, rue de St Brieue, 35042 Rennes Cedex Translator: Dr. Jenny Watson, Department of Zoology, University of Aberdeen. Translation Editor: Dr. Lindsay Laird, Department of Zoology, University of Aberdeen. Original French edition, La Truite: biologie et ecologie Published by © INRA, Paris, 1991 This work has been published with the help of the French Ministere de la Culture. SPRINGER-PRAXIS SERIES IN AQUACULTURE & FISHERIES SERIES EDITOR Dr. Lindsay Laird, M.A., Ph.D., University of Aberdeen, UK CONSULTANT EDITOR Dr. Selina Stead, B.Sc., M.Sc., Ph.D., Scottish Agricultural College, Aberdeen, UK ISBN 978-1-85233·317·1 e-ISBN-13 :978·1·4471·0775-0 DOl: 10.10071978·1·4471·0775·0 British Library Cataloging in Publication Data Bagliniere, I. L. Biology and ecology of the brown and sea trout. . (Springer-Praxis series in aquaculture and fisheries) l.Sea·run brown trout 2.Sea-run brown trout -Ecology I.Title II.Maisse, G. 597.5'7 ISBN-13: 978-1-85233-317-1 Library of Congress Cataloging-in-Publication Data Truite, biologie et ecologie. English Biology and ecology of the brown sea troutl[edited by) I.L. Bagliniere, G. Maisse: [translator, Jenny Watson]. p. cm. --(Spring-Praxis series in acquaculture & fisheries) ISBN-13:978-1-85233-317-1 (alk. paper) I. Brown trout. l. Bagliniere, Jean-Luc. II. Maise, G. III. Title. IV. Series. QL638.S2T8213 1999 597.5'7 --dc21 98-51613 CIP Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the Copyright, Designs and Patents Act 1988, this publication may only be reproduced, stored or transmitted, in any form or by any means, with the prior permission in writing of the publishers, or in the case of repro graphic reproduction in accordance with the terms oflicences issued by the Copyright Licensing Agency. Enquiries concerning reproduction outside those terms should be sent to the publishers. © Praxis Publishing Ltd, Chichester, UK, 1999 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Cover design: lim Wilkie Typesetting: Heather FitzGibbon, Christchurch, Dorset, UK Printed on paper supplied by Precision Publishing Papers Ltd, UK Contents Acknowledgements .................................................. vii Introduction: The brown trout (Salmo trutta L.): its origin, distribution and economic and scientific significance J. L Bagliniere ................................................ . Part I. River trout 1. Biology of the brown trout (Salmo trutta L.) in French rivers G. Maisse and J. L. Bagliniere .................................... 15 2. The habitat of the brown trout (Salmo trutta L.) in water courses J. Haury, Dominique Ombredane and J. L Bagliniere . . . . . . . . . . . . . . . . . . 37 3. Feeding strategy of the brown trout (Salmo trutta L.) in running water A. Neveu...................................................... 91 4. Social organization and territoriality in brown trout juveniles during ontogeny M. Heland ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 115 Part II. Ecological plasticity and genetic diversity in trout l. Main characteristics ofthe biology of the trout (Salmo trutta L.) in Lake Leman (Lake Geneva) and some of its tributaries A. Champigneulle, B. Buttiker, P. Durand and M. Melhaoui ............ 147 2. Sea trout (Salmo trutta L.) in Normandy and Picardy G. Euzenat, Fran~oise Fournel and A. Richard. . . . . . . . . . . . . . . . . . . . . .. 175 3. Genetic diversity and the management of natural populations of brown trout R. Guyomard . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 205 vi Contents Part Ill. The management of natural populations of brown trout The management of natural populations of brown trout in France, analysed from an historical perspective (1669-1986) M. Thibault . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 227 Conclusion: Current ideas on the biological basis for management of trout populations (Salmo trutta L.) G. Maisse and J. L. Bagliniere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 277 Index ............................................................. 283 The colour illustration of brown trout habitats is facing page 42. Acknowledgements This book was conceived by the scientific committee of the Colloquium on Trout, organ ized from 6 to 8 September 1988 at the Centre du Paraclet (Conseil Superieur de la Peche) by INRA, the National Agronomic Research Institute (Physiology and Ecology of Fishes Station) and the Conseil Superieur de la Peche (French Bulletin of Fishing and Fish Culture). The colloquium was financed by SPRETIE (the Secretariat de l'Etat a l'Environnement), the Conseil Superior de la Peche and INRA. Other articles presented during this colloquium have been published in two editions of the Bulletin Fran~ais de Peche et de Pisciculture (nos 318 and 319 and special Trout Colloquium edition). The scientific committee consisted of: J. Allardi (CEMAGREF, Paris), J. Arrignon (Union Nationale des Federations de Peche), J. L. Bagliniere (INRA, Rennes), B. Buttiker (Conservatoire de la Faune, Switzerland), A. Champigneulle (INRA, Thonon-Ies-Bains), B. Chevassus (INRA, Paris), Y. Cote (Minister for Leisure, Hunting and Fishing, Quebec), P. Dumont (Minister for Leisure, Hunting and Fishing), Fran~oise Fournel (CSP, Compiegne), M. Heland (INRA, St P6eINivelle), G. Maisse (INRA, Rennes), A. Neveu (INRA, Rennes), A. Nihouarn (CSP, Rennes), J. C. Phillipart (University of Liege, Belgium), A Richard (CSP, Rennes), E. Vigneux (CSP, Paraclet). We wish to thank all the people who, through their remarks and suggestions, helped to improve the quality of the articles presented in this book: J. Arrignon (Union Nationale des Federations de Peche), P. Bergot (INRA, St P6elNivelle), B. Chevassus (INRA, Jouy en-Josas), Brigitte Dasaigues (University of Paris I), P. Gaudin (University of Claude Bernard Lyon I), J. Y. Gautier (University of Rennes), J. Genermont (University of Paris South), D. Gerdeaux (INRA, Thonon-Ies-Bains), R. Guyomard (INRA, Jouy-en-Josas), M. Heland (INRA, St P6elNivelle), C. Lagier (University of Lyon IT), P. Y. Lebail (INRA, Rennes), J. Lecomte (INRA, Jouy-en-Josas), A. Neveu (INRA, Rennes), Dominique Ombredane (INRA, Rennes), E. Prevost (INRA, Rennes), M. Thibault (INRA, Rennes). Introduction: The brown trout (Salmo trutta L)-its origin, distribution and economic and scientific significance 1. L. Bagliniere I. INTRODUCTION The brown trout is a species of salmonid with a facultative migratory character (Hoar, 1976) and has a great capacity for adaptation to different environments. This has led to a major degree of polymorphism in this species, which has been classified in the past under different scientific names (Melhaoui, 1985; Elliott, 1989, 1994). However, the interpreta tion of the polymorphy of the trout remains delicate in terms of genetic differences (Krieg, 1984), but the idea of the existence of a single species, Salmo trutta Linnaeus, actually remains the most likely. n.PHYLOGENY The genus Salmo constitutes, with six others (Brachymystax, Salmothymus, Acantholinqua, Hucho, Salvelinus, and Oncorhynchus), the sub-family Salmonines, one of three making up the Salmonid family (Fig. 1) (Nelson, 1994). This genus comprises only two species, Salmo trutta, the brown trout and Salmo salar the Atlantic salmon, since the rainbow trout (or steelhead), Salmo gairdneri and the cut-throat, Salmo clarkii have recently been reclassified in the genus Oncorhynchus (Smith and Stearley, 1989). But the systematics of the genus Salmo is still not quite clear as some subspecies of the brown trout can be considered as species of the genus Salmo according to author (i.e. Salmo (Trutta) carpio and Salmo (Trutta) marmoratus (see Fig. 2) (Elliott, 1994; Berrebi, 1997). The ancestors of the Salmonid family appeared at the start of the Cretaceous period (between 63 and 135 million years ago) (Legendre, 1980). According to Tchemavin (1939), they would originate in freshwater. The sub-family Salmoninae appeared after the tertiary period, more accurately the Miocene (between 13 and 25 million years ago) J. L. Baglinière et al. (eds.), Biology and Ecology of the Brown and Sea Trout © Praxis Publishing Ltd, Chichester, UK 1999 2 Introduction ORDER Salmoniformes j FAMILY Salmomdae SUB-F AMIL Y Coregonidae Salmoninae Thymallinae GENUS Oncorhynchus Salmo Salvelinus SPECIES Salmo trutta Salmo salar Fig. 1. (Legendre, 1980). The three main genera (Salmo, Oncorhynchus, Salvelinus) known to day appeared more recently (at the start of the Pleistocene) from this common ancestry (Jones, 1959). The separation of the American and Euro-Asiatic continents and the subse quent succession of glaciation ages of the Pleistocene and recent eras has not only resulted in the differentiation of these three genera but also the appearances of numerous lower level taxa (Jones, 1959; Hoar, 1976). However, according to Tchernavin (1939), anadromous behaviour appeared at the start of the Ice Age. However, Balon (1980) considers that this behaviour existed before speciation in the Salmonid family and that the marine types, mainly of the genus Salmo would have formed the origin of the freshwater forms. From this well-argued theory of Balon (1980), Thorpe (1982) deduced that the salmonids were in fact primitive teleosts, probably of marine origin, and that certain species may have progressively lost their anadromous behaviour. The formation of the sub-family Salmonines occurred at the end of the last glaciation period, about 10 000 years ago. It was linked to temporary geographical barriers caused by the advancing or retreating glaciers and to temperature changes in the oceans (Jones, 1959). In the case of the trout, Salmo trutta, the appearance of the anadromous type at the time of the glaciations provided the origin of a large variety of freshwater European forms (Lelek, 1980) (Fig. 2). ill. DISTRIBUTION 1. Original distribution (Fig. 2) The brown trout is essentially present, in its many forms, across Europe. The northern most limits of its distribution stretch from Iceland to the USSR (north of the Volga), m --' 500 k----- ' ' P : ~cN EA 2. RAN Fig. R E T DI E M TH A R E O S N -"-.. ~..... \ ,.' i 1 +2 If " '\ '", •• , TLANTIC OCEAN A