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BIOLOGY AND ECOLOGY OF PIKE BIOLOGY AND ECOLOGY OF PIKE Editors Christian Skov DTU Aqua, Technical University of Denmark Section for Inland Fisheries and Ecology Silkeborg, Denmark P. Anders Nilsson Lund University Department of Biology – Aquatic Ecology Lund, Sweden and Karlstad University Department of Environmental and Life Sciences – Biology Karlstad, Sweden Cover illustration: Reproduced by kind courtesy of Olof Engstedt (author of Chapter 10). CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 © 2018 by Taylor & Francis Group, LLC CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works Printed on acid-free paper Version Date: 20171023 International Standard Book Number-13: 978-1-4822-6290-2 (Hardback) This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the consequences of their use. The authors and publishers have attempted to trace the copyright holders of all material reproduced in this publication and apologize to copyright holders if permission to publish in this form has not been obtained. If any copyright material has not been acknowledged please write and let us know so we may rectify in any future reprint. Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying, microfilming, and recording, or in any information storage or retrieval system, without written permission from the publishers. For permission to photocopy or use material electronically from this work, please access www.copyright.com (http://www.copyright.com/) or contact the Copyright Clearance Center, Inc. (CCC), 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. CCC is a not-for-profit organization that provides licenses and registration for a variety of users. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Library of Congress Cataloging-in-Publication Data Names: Skov, Christian, 1968- editor. | Nilsson, P. Anders, editor. Title: Biology and ecology of pike / editors, Christian Skov, DTU Aqua, Technical University of Denmark, Section for Inland Fisheries and Ecology, Silkeborg, Denmark, P. Anders Nilsson, Lund University, Department of Biology & Aquatic Ecology, Lund, Sweden, and Karlstad University, Department of Environmental and Life Sciences & Biology, Karlstad, Sweden. Description: Boca Raton, FL : CRC Press, 2017. | “A Science Publishers book.” | Includes bibliographical references and index. Identifiers: LCCN 2017034002 | ISBN 9781482262902 (hardback : alk. paper) Subjects: LCSH: Pike. | Pike fisheries--Management. Classification: LCC QL638.E7 B56 2017 | DDC 597.5/9--dc23 LC record available at https://lccn.loc.gov/2017034002 Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com To the ladies of our lives Camilla, Felicia, Hanna, Dorte, Liv. And to Alexander. Contents P. Anders Nilsson1 and Peter Eklöv2 1. Preface; Introduction to Pike and This Book 1 P. Anders Nilsson and Christian Skov Section I Individuals 2. Finding Food and Staying Alive 9 P. Anders Nilsson and Peter Eklöv 3. Coping with Environments; Vegetation, Turbidity and Abiotics 32 Lene Jacobsen and Jonna Engström-Öst 4. Bioenergetics and Individual Growth of Pike 62 John D. Armstrong 5. Spatial Ecology 83 Christian Skov, Martyn C. Lucas and Lene Jacobsen Section II Populations and Communities 6. Pike Population Size and Structure: Influence of Density-Dependent and Density-Independent Factors 123 Thrond O. Haugen and L. Asbjørn Vøllestad 7. Population Genetics of Pike 164 Lovisa Wennerström, Dorte Bekkevold and Linda Laikre 8. Trophic Interactions 185 Anders Persson, P. Anders Nilsson and Christer Brönmark viii Biology and Ecology of Pike Section III Management and Fisheries 9. Stocking for Pike Population Enhancement 215 Nicolas Guillerault, Daniel Hühn, Julien Cucherousset, Robert Arlinghaus and Christian Skov 10. Habitat Restoration – A Sustainable Key to Management 248 Olof Engstedt, Jonas Nilsson and Per Larsson 11. Pike Stocking for Lake Restoration 269 Christian Skov 12. Recreational Piking - Sustainably Managing Pike in Recreational Fisheries 288 Robert Arlinghaus, Josep Alós, Ben Beardmore, Ángela M. Díaz, Daniel Hühn, Fiona Johnston, Thomas Klefoth, Anna Kuparinen, Shuichi Matsumura, Thilo Pagel, Tonio Pieterek and Carsten Riepe 13. Northern Pike Commercial Fisheries, Stock Assessment and Aquaculture 337 Anna Kuparinen and Hannu Lehtonen 14. The Northern Pike, A Prized Native but Disastrous Invasive 356 Kristine Dunker, Adam Sepulveda, Robert Massengill and David Rutz Index 399 Chapter 1 Preface; Introduction to Pike and This Book P. Anders Nilsson*1 and Christian Skov2 The pike (Esox lucius Linnaeus, 1758), or northern pike as it is called in most of North America, is an iconic fish species that has fascinated mankind throughout history. Its large size and characteristic appearance have likely contributed to various tall stories, some of which are beautifully saved in Izaak Walton’s “The Compleat Angler or the Contemplative man’s Recreation” from 1653 (see e.g. Walton and Cotton 2005). The pike is there described as the tyrant of freshwaters, attacking everything from venomous frogs to birds to mules and people. There is also reference to anecdotes of pike being spontaneously generated from weed and glutinous matter with help of sunlight, and of specimens over 200 years old. Although such historical texts probably contain various degrees of misconception, they simultaneously nicely describe many aspects of pike as a fish species and fisheries target, and, perhaps most importantly, indicate how significant pike as a species has been and continues to be. Along similar lines, the pike has undoubtedly interested us for many years, from both personal and academic points of view, and it is with the abovementioned importance, significance and fascination in mind we have set the current book. The last century shows an increasing and continuing interest in pike in the scientific literature (Fig. 1.1), warranting our summary of pike knowledge. Needless to say, it is impossible to cover all of those publications and knowledge *Corresponding author: 1Lund University, Department of Biology – Aquatic ecology, Ecology Building, 22362 Lund, Sweden; Email: [email protected] and Karlstad University, Department of Environmental and Life Sciences – Biology, 65188 Karlstad, Sweden; Email: [email protected] 2DTU Aqua, Technical University of Denmark, Section for Inland Fisheries and Ecology, Vejlsøvej 39 8600, Silkeborg, Denmark; Email: [email protected] 2 Biology and Ecology of Pike on pike in a single book. Also, the scientific interest in and knowledge of pike have previously been reviewed in synopses, books and journal special issues, such as Raat (1988), Craig (1996), Pierce (2012) and Farrell et al. (2008). Although these volumes remain substantially influential to the pike literature and knowledge, they leave temporal and geographical knowledge gaps to the understanding of pike. To complement and extend from these volumes, as well as to update the current state of pike research and knowledge, the current contribution focuses on pike from three main angles in three sections; pike individual ecology, pike populations and their communities, as well as management and fisheries of pike. FIGURE 1.1 The annual (1916-2016) number of scientific publications including the search term “Esox lucius”, extracted from The Web of Science. Adaptive decisions are made by individuals, and as such decisions can affect, for example, pike individual success, growth and survival, they have bearing also for pike population densities and size structures, as well as for trophic processes and management strategies. This volume therefore starts with a section on pike Individuals, with chapters focusing on pike foraging behaviour and decisions, how pike cope with different environments, the bioenergetics and growth of pike, and pike spatial ecology. In chapter 2, pike foraging behaviour is described and scrutinized from a predation-cycle perspective, highlighting effects of pike and prey interactions, as well as pike risks of predation, kleptoparasitism and cannibalism, on pike foraging performance, prey preference, and functional responses. Pike individual success moreover depends on the environment they live in, and chapter 3 deals with crucial aspects of vegetation, water clarity, temperature, oxygen, salinity, and pH for different pike life stages, all with possible climate change in mind. Preface; Introduction to Pike and This Book 3 The consequences of foraging and environment should be translated into growth and reproduction to contribute to lifetime fitness, and pike bioenergetics, growth and physiological capacities are considered in chapter 4, in light of pike energy budgets, emphasising the special adaptations pike possess. The last chapter of the Individuals section of the book deals with the
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