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Ecological Studies, Vol. 135

Analysis and Synthesis

Edited by

M. M. Caldwell, Logan, USA

G. Heldmaier, Marburg, Germany

0. L. Lange, Wiirzburg, Germany

H. A. Mooney, Stanford, USA

E.-D. Schulze, Bayreuth, Germany

U. Sommer, Kiel, Germany

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Ecological Studies

Volumes published since 1992 are listed at the end of this book.

Springer-Verlag Berlin Heidelberg GmbH

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Predation

in Vertebrate Communities

The Bialowieia Primeval Forest as a Case Study

With 251 Figures and 103 Tables

~ Springer

B. Jędrzejewska W. Jędrzejewski

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Dr. BoGUMILA J:~DRZEJEWSKA

Dr. Wt.ODZIMIERZ J:~DRZEJEWSKI

Mammal Research Institute Polish Academy of Sciences Ul. Gen. Waszkiewicza 1 17-230 Bialowieza, Poland

ISSN 0070-8356 ISBN 978-3-642-08384-6

Library of Congress Cataloging-in-Publication Data

J~drzejewska, B. (Bogumita), 1959- . Predation in vertebrate communities: the Bialowieza Primeval Forest as a case study I B. j~drzejewska, W. J~drzejewski. p. em. - (Ecological studies; vol. 135) Includes bibliographical references (p. ) and index.

ISBN 978-3-642-08384-6 ISBN 978-3-662-35364-6 (eBook) DOI 10.1007/978-3-662-35364-6

I. Predation (Biology) - Bialowieza Forest (Poland and Russia) 2. Population biology - Bialowieza Forest (Poland and Russia) 3. Vertebrates - Ecology - Bialowieza Forest (Poland and Russia) I. )~drzejewski, W.

(Wlodzimierz), 1961- . II. Title. Ill. Series: Ecological studies; v. 135. QL758.)44 1998 596.153-dc21 98-2772 CIP

This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcast- ing, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this pub- lication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag Berlin Heidelberg GmbH.

Violations are liable for prosecution under the German Copyright Law.

©Springer-Verlag Berlin Heidelberg 1998

Originally published by Springer-Verlag Berlin Heidelberg New York Tokyo in 1998 Softcover reprint of the hardcover 1st edition 1998

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 protec- tive laws and regulations and therefore free for general use.

Production: PRO EDIT GmbH, D-69126 Heidelberg

Cover design: Design & Production GmbH, D-69121 Heidelberg Typesetting: K+V Fotosatz GmbH, D-64743 Beerfelden

SPIN 10482555 31/3137-5 4 3 2 l 0 - Printed on acid-free paper

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Acknowledgements

Our work on this book and the 10-year study on Bialowieza's predators were, in some ways, the result of team work, and it is a pleasure to acknowledge the help of many friends, colleagues, and teachers. Jacek Goszczynski, Joan- na Gliwicz, and Marek Keller, our professors at the Forestry Faculty of War- saw Agricultural University, fostered our scientific interest in vertebrate ecol- ogy and - for years after we had graduated - encouraged and helped us through advice, constructive criticism and discussions. We are grateful to Professor Zdzislaw Pucek, the Director of the Mammal Research Institute (Polish Academy of Sciences) in Bialowieza, for his unfailing support for our wide-scale research programme, scrupulous reading and correcting of our scientific texts, and encouragement during the work on this book.

Since 1985, when we began the investigations, the research couple has grown into a research team, the Group of Predator Ecology, a group of en- thusiasts. Many people who joined us, for shorter or longer periods of coop- eration, have contributed enormously to the data synthesised in this book.

We thank our colleagues-researchers, Andrzej Zalewski, Krzysztof Schmidt, Henryk Okarma, Rafal Kowalczyk, and Leszek Rychlik, and graduate stu- dents, Marcin Brzezinski, Wojciech Nowakowski, Alicja Morylewska, Janusz Kupis, and Marco Musiani. We are very grateful to the technicians who worked on our projects in 1985-1996: Irena Szpakowicz, Nadzieja Wojcie- szuk, Swietlana Bogdanska, Alicja Buszko, Edyta Bajko, Jerzy Siemieniuk, Miroslaw Szlachciuk, Marek Swic, Eugeniusz Bujko, Stefan Buszko, Maksymi- lian Szuma, Arkadiusz Arasim, Stanislaw Sniezko, and the late Leonidas Sie- mieniuk. Their jobs included caring for laboratory rodents and predators, snow tracking, live trapping of animals, radio-tracking, laboratory analysis of scats and pellets, computer work and much more. We are particularly in- debted to Lucyna Szymura and Karol Zub for their tremendous work. Lucy- na put most of the data into computer files, helped in organising field cen- suses, and prepared microscope slides of mammalian hair recovered from predator scats and pellets. Karol identified a large part of the prey remnants (especially birds, amphibians, and insects), analysed graphically the intricate data on space use by radio-tracked carnivores, and prepared infinite num- bers of figures in the computer programme CorelDraw. We thank them for a creative attitude to all tasks they were fulfilling. We are grateful to all other workers of the Mammal Research Institute, who helped us in many ways.

We appreciate the financial and technical assistance from Earthwatch (USA) and the work of Earth Corps volunteers on our project. Thanks are extended to Linda Wallace-Frjedland, Bill Smith, Marylouise Morgan (USA), Don and Betty

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VI Acknowledgements

Wood (Australia) and other people for helping us in various ways. We also thank many other volunteers from Poland and other countries, for whom no fieldwork was too hard and no time spent in the forest was too long.

The development of our long-term study on predators in Bialowieia Forest brought us closer to other investigators and fostered very useful and satisfacto- ry cooperation with Drs. Andrzej L. Ruprecht, Malgorzata Krasinska (Mammal Research Institute), Zbigniew A. Krasinski (Bialowieia National Park), Marcin Brzezinski (Warsaw University), Lech Milkowski (Bialowieia State Forestry District), Arkadiusz Szymura (Forestry High School, Bialowieia), Eugeniusz Pugacewicz (North-Podlasie Society for Bird Protection), Aleksei N. Bunevich (State National Park Belovezhskaya Pushcha, Belarus Republic), Vadim E. Si- dorovich and Mikhail M. Pikulik (Institute of Zoology, Belarussian Academy of Sciences, Minsk). We thank them all for inspiration and sharing their experi- ence and knowledge with us.

Several zoologists have made their yet unpublished findings available for in- corporation in this book. They include Drs. Zdzislaw Pucek, Marcin Brzezinski, Vladimir A. Dackevich, Aleksei N. Bunevich, Lech Milkowski, Vadim E. Sidor- ovich, Mikhail M. Pikulik, and Wieslaw Walankiewicz. To all these people we are grateful for their help. We thank Lech Z. Nowacki, the author of line draw- ings, and the photographers whose pictures of wildlife and Bialowieia Forest enliven this book.

The birth and growth of many ideas presented in this book were stimulat- ed by numerous discussions - in personal contacts and letters - with Drs.

Zdzislaw Pucek, Jacek Goszczynski, Joanna Gliwicz, Ludwik Tomialojc, To- masz Wesolowski, Wieslaw Walankiewicz (Poland), Burton L. Gordon, George Batzli, (USA), Lennart Hansson, Sam Erlinge (Sweden), Ilkka Hanski, Hannu Ylonen, Erkki Korpimaki (Finland), Eivind Ostbye, Nils C. Stenseth, Geir A. Sonerud (Norway), and Carolyn King (New Zealand). Drs. Mikael Sandell (Sweden), Urs Breitenmoser (Switzerland), and L. David Mech (USA) taught us how to capture and radiotrack carnivores.

Finally, we express our gratitude to managers and employees of the following institutions that permitted our access to the study area and helped in various ways: Bialowieia National Park (Dr. Czeslaw Okol6w, Director, and Prof. Ja- nusz B. Falinski,. Chairman of the Scientific Council), State National Park Belo- vezhskaya Pushcha (Mr. Vasilii P. Zhukov, General Director, Dr. Vyacheslav V.

Semakov, Director of the Science Department), State Forestry Districts of Bialowieia (Mr. Stanislaw Ignatiew), Hajn6wka (Mr. J6zef Fularczyk), and Browsk (Mr. Andrzej Jaworski), and the Forestry High School in Bialowieia.

The work on this book was financed by the grant KBN 4 4416 91 02 from the Committee for Scientific Research (Poland) and the budget of the Mam- mal Research Institute.

Many individuals and institutions contributed to the good points in this book. For the weak points, the responsibility is ours.

Bialowie:ia, April 1998 B. JJ?DRZEJEWSKA

W. JJ?DRZEJEWSKI

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Contents

Introduction . . . 1

1 Bialowieza Primeval Forest - Abiotic Conditions and Habitats . . 4

2 Abundance of Prey . . . 28

2.1 Introduction . . . 28

2.2 Methodical Outline . . . 28

2.3 Ungulates - A Royal Feast Served on Too High Tables . . . 32

2.4 Rodents - Voles and Mice Everywhere . . . 48

2.5 Brown Hare . . . 58

2.6 Shrews and Moles . . . 58

2.7 Bats - Uncatchable Flying Objects . . . 66

2.8 Birds - Endless Choice of (Expensive) Meals to Predators . . . 67

2.9 Reptiles - Rare Animals of Unpredictable Activity . . . 86

2.10 Frogs - Plentiful in Rainy Summers, None in Winters . . . 88

2.11 Vertebrate Herbivores and Insectivores of Temperate Forests - Characteristics of Prey Resources for Carnivores and Raptors . . . 92

3 Numbers, Distribution and Home Ranges of Predators . . . 97

3.1 Introduction . . . 97

3.2 An Outline of Methods Used to Survey Predators . . . 98

3.3 Lynx (Lynx lynx) . . . 102

3.4 Wolf (Canis lupus) ... 106

3.5 Red Fox (Vulpes vulpes) ... 111

3.6 Raccoon Dog (Nyctereutes procyonoides) ... 114

3.7 Badger (Meles meles) . . . 115

3.8 Pine Marten (Martes martes) ... 117

3.9 Stoat (Mustela erminea) . . . 121

3.10 Weasel (Mustela nivalis) ... 121

3.11 Polecat (Mustela putorius) ... 125

3.12 River Otter (Lutra lutra) ... 128

3.13 American Mink (Mustela vison) . . . 129

3.14 Goshawk (Accipiter gentilis) ... 131

3.15 Sparrowhawk (Accipiter nisus) ... 134

3.16 Common Buzzard (Buteo buteo) ... 135

3.17 Rough-Legged Buzzard (Buteo lagopus) ... 138

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VIII Contents

3.18 Honey Buzzard (Pernis apivorus) ... 139

3.19 Lesser Spotted Eagle (Aquila pomarina) ... 141

3.20 Large and Rare Eagles: Golden Eagle (Aquila chrysaetos), Short-Toed Eagle (Circaetus gallicus), White-Tailed Eagle (Haliaeetus albicilla), and Booted Eagle (Hieraaetus pennatus) ... 143

3.21 Black Kite (Milvus migrans) and Red Kite (Milvus milvus) ... 145

3.22 Marsh Harrier (Circus aeruginosus) and Montagu's Harrier (Circus pygargus) . . . 146

3.23 Hobby (Falco subbuteo) ... 147

3.24 Tawny Owl (Strix aluco) ... 148

3.25 Pygmy Owl (Glaucidium passerinum) ... 153

3.26 Tengmalm's Owl (Aegolius funereus) ... 155

3.27 Long-Eared Owl (Asio otus) . . . 156

3.28 Large Owls: Eagle Owl (Bubo bubo) and Great Grey Owl (Strix nebulosa) ... 158

3.29 Predators No Longer Breeding in Bialowie:ia Primeval Forest and Records of Non-Breeding Raptors ... 159

3.30 Predators Inhabiting Human Settlements . . . 161

3.31 Community of Vertebrate Predators - Species Structure, Densities, and Habitat Distribution ... 162

3.32 Factors Shaping Densities and Population Dynamics of Predators 173 4 Diets and Hunting Habits of Predators ... 180

4.1 Introduction ... 180

4.2 Methods of Studying Predator Feeding Habits ... 181

4.3 Lynxes - Stalking the Roe Deer ... 186

4.4 Wolves- Running After Red Deer ... 198

4.5 Red Foxes - Hunting for Voles in Meadows and Scavenging in Woodland . . . 208

4.6 Raccoon Dogs- Scavenging on Ungulate Carcasses ... 215

4.7 Badgers and Earthworms ... 219

4.8 Pine Martens - Generalistic Feeding Habits of Bank Vole Specialists . . . 224

4.9 Stoats - Digging for Rodents . . . 231

4.10 Weasels - Rodent Specialists ... 233

4.11 Polecats' Cold Meals of Frogs . . . 23 7 4.12 River Otters - Between Fishing and 'Frogging' . . . 240

4.13 American Minks- Hunting Fish and Frogs in Summer, Frogs and Rodents in Winter . . . 245

4.14 Goshawks- Year-Round Pursuit of Birds ... 249

4.15 Sparrowhawks - Raptors Specialised on Small Forest Birds ... 254

4.16 Buzzards - Shifting from Voles to Birds and Moles ... 258

4.17 Honey Buzzards- A Misleading Name for Wasp-Eaters ... 266

4.18 Lesser Spotted Eagles - Shifting from Amphibians to Birds, to Mammals . . . 268

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Contents IX

4.19 Short-Toed Eagles - Specialisation on Snakes

and Snake-Like Lizards ... 275

4.20 Booted Eagles - Hunting on Medium-Sized Birds of Open Land and Forests ... 275

4.21 Black Kites - 'River Ospreys' of Temperate Forests, and Red Kites- Unspecialised Hunters and Scavengers ... 278

4.22 Hobbies - Aerial Hunting for Swallows, Larks, and Dragonflies . . 280

4.23 Tawny Owls- Specialisation on Yellow-Necked Mice with Secondary Adaptations for Hunting Birds, Shrews, and Frogs 284 4.24 Pygmy Owls- Tyrants of Passerine Birds ... 291

4.25 Tengmalm's Owls - Boreal Menu of Bank Voles, Tits, and Goldcrests . . . 295

4.26 Long-Eared Owls - Specialisation on Microtus Voles ... 295

4.27 Eagle Owls - Night Hunters of Large Birds . . . 295

4.28 Peregrines - Sky Diving for Large Birds . . . 297

4.29 Food of Predators Inhabiting Human Settlements: Barn Owls and Stray Dogs . . . 299

4.30 Searching Modes of Carnivores in the Pristine Temperate Forest 301 4.31 Which Prey Groups Matter Most in the Diets of Temperate Forest Predators? . . . 304

4.32 The Pattern of Sharing Resources - Food Niche Overlaps . . . 308

4.33 Breadth of Food Niches and Choice of Primary and Secondary Prey ... 312

4.34 Foraging Strategies and Functional Responses of Predators 320 5 Predation Impact and Its Importance to Prey Populations 324 5.1 Introduction ... 324

5.2 Methods of Calculating the Predation Impact . . . 328

5.3 Total Consumption of Prey Biomass by All Predators . . . 331

5.4 Predation on Ungulates . . . 335

5.4.1 Species Structure of Total Predation on Ungulates ... 335

5.4.2 Magnitude and Mechanisms of Wolf and Lynx Predation on Ungulates . . . 336

5.4.3 Relative Roles of Predation and Other Factors in Shaping the Densities and Population Dynamics of Ungulates 347 5.5 Predation on Rodents . . . 358

5.5.1 Species Structure of Total Predation on Rodents . . . 358

5.5.2 Magnitude and Mechanisms of Predation on Forest Rodents . . . . 360

5.5.3 Relative Roles of Predation and Other Factors in Shaping the Densities and Population Dynamics of Rodents . . 367

5.5.4 Antipredatory Behaviours of Forest Rodents ... 374

5.6 Predation on Shrews . . . 375

5.6.1 Species Structure of Total Predation on Insectivores ... 375

5.6.2 Magnitude and Mechanisms of Predation on Shrews . . . 377

5.7 Predation on Birds ... 380

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X Contents

5.7.1 Species Structure of Total Predation on Birds ... 380

5.7.2 Magnitude and Mechanisms of Predation on Birds ... 382

5.7.3 Relative Roles of Predation and Other Factors in Shaping the Densities and Population Dynamics of Bird . . . 389

5.8 Predation on Amphibians ... 401

5.8.1 Species Structure of Total Predation on Amphibians ... 401

5.8.2 Magnitude and Mechanisms of Predation on Amphibians . . . 403

5.8.3 Roles of Predation and Other Factors in Shaping the Densities and Population Dynamics of Amphibians . . . 405

5.9 Intraguild Predation in Carnivores and Raptors of Bialowieza Forest . . . 408

5.10 Scavenging- Seemingly a 'Free Lunch', Sometimes at High Cost . 411 5.10.1 Sources and Supply of Ungulate Carcasses ... 411

5.10.2 Utilisation of Carcasses by Scavengers ... 415

5.11 Predation in Vertebrate Animal Communities of the European Temperate Forests- An Attempt at Synthesis ... 421

References ... 427

Subject Index . . . 443

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