Tài liệu Carnivore Ecology and Conservation A Handbook of Techniques pptx

Carnivore Ecology and Conservation

OUP CORRECTED PROOF – FINAL, 6/12/2011, SPi

Techniques in Ecology and Conservation Series

Series Editor: William J. Sutherland

Bird Ecology and Conservation: A Handbook of Techniques

William J. Sutherland, Ian Newton, and Rhys E. Green

Conservation Education and Outreach Techniques

Susan K. Jacobson, Mallory D. McDuff, and Martha C. Monroe

Forest Ecology and Conservation: A Handbook of Techniques

Adrian C. Newton

Habitat Management for Conservation: A Handbook of Techniques

Malcolm Ausden

Conservation and Sustainable Use: A Handbook of Techniques

E.J. Milner-Gulland and J. Marcus Rowcliffe

Invasive Species Management: A Handbook of Principles and Techniques

Mick N. Clout and Peter A. Williams

Amphibian Ecology and Conservation: A Handbook of Techniques

C. Kenneth Dodd, Jr.

Insect Conservation: A Handbook of Approaches and Methods

Michael J. Samways, Melodie A. McGeoch, and Tim R. New

Remote Sensing for Ecology and Conservation: A Handbook of Techniques

Ned Horning, Julie A. Robinson, Eleanor J. Sterling, Woody Turner, and Sacha Spector

Marine Mammal Ecology and Conservation: A Handbook of Techniques

Ian L. Boyd, W. Don Bowen, and Sara J. Iverson

Carnivore Ecology and Conservation: A Handbook of Techniques

Luigi Boitani and Roger A. Powell

OUP CORRECTED PROOF – FINAL, 6/12/2011, SPi

Carnivore Ecology and

Conservation

A Handbook of Techniques

Edited by

Luigi Boitani

and

Roger A. Powell

1

OUP CORRECTED PROOF – FINAL, 6/12/2011, SPi

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Foreword

Animals that must hunt and kill for at least part of their living are inherently

interesting to many people. Perhaps that is because humans evolved to make our

living that way as well, and carnivores often compete with us to this very day.

Wolves, bears, lions, tigers, leopards, lynx, mink, weasels, and foxes, and a wide

variety of their relatives, have long grabbed the human imagination. In any case,

carnivores comprise a very significant contingent of the world’s wildlife, and many

books have been written about them.

This book is distinct from its predecessors primarily through its emphasis on

techniques for dealing with carnivores: how to sample them, capture them for study,

handle them, monitor them, and even how to help minimize their competition with

us. It is a very helpful book that fills an important niche and comes at the right time.

In many parts of the world carnivores are persecuted, while in other parts they

are being restored. Thus societies remain interested in carnivores for one reason or

another, and science serves society’s interest through numerous carnivore studies.

The authors of this book’s chapters have conducted a significant proportion of

those studies for many years, and the editors for even longer.

Both editors are well qualified to produce this book, having studied and worked

with carnivores and their conservation for decades. I had the great opportunity of

partnering with Luigi Boitani in 1974, early in his career, when we spent a month in

Italy’s Abruzzo Mountains live-trapping, radio-collaring, and tracking wolves. I had

presented my paper “Current Techniques in the Study of Elusive Wilderness

Carnivores” at the Eleventh International Congress of Game Biologists in Stock￾holm in September 1973. It covered my experiences live-trapping and radio￾tracking wolves, fishers, martens, and lynxes as well as a literature review of current

techniques used to study other carnivores. I like to think of that paper as a germ that

helped spawn the present book. Luigi attended the Stockholm meeting, sought to

apply my techniques with wolves in Italy, and asked me to join him there to get

started. I eagerly agreed. Little did I realize then that 40 years later, Luigi and Roger

Powell would devote a whole book to techniques for studying carnivores.

During the same general period when I met Luigi, I also met Roger Powell.

Roger had joined my research team as a summer intern on a wolf–deer project in

the Superior National Forest of Minnesota, where we had also been radio-tracking

lynx, martens, and fishers on the side. The duties clearly agreed with him, for a few

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years later he began his own carnivore study, this one involving fishers. That study

became his dissertation topic, and I became one of his advisors.

That was all long ago, and the field has advanced greatly and blossomed. Now

instead of merely locating an animal via telemetry (a feat in itself years ago), one

searches the profuse literature, decides on study objectives, carefully plans the

study’s design, and chooses from any of the many high-tech radio-collars on the

commercial market that will best serve the objectives.

However, dealing with the most appropriate technology to study carnivores is

only a small part of carnivore investigations now. The data currently obtainable has

opened many new carnivore research vistas, and Boitani and Powell and their

collaborators have assembled a set of chapters that nicely address that array. An

early chapter on carnivore surveys, for example, is basic, for such surveys are of

special importance, both spatially and temporally. In some areas and with some

species, just obtaining a general idea of numbers and distribution can be very

important. Mapping such distributions plays a major role in these studies, and

non-invasive sampling is particularly valuable, especially with endangered or rare

species and in inaccessible areas. These subjects are well covered in this book.

In some areas of the world and with certain carnivores, detailed counts are

required annually. Sometimes with such counts it is valuable to estimate various

demographic parameters, and radio-telemetry often facilitates those estimates. To

collar carnivores, it is necessary to capture and handle them, allowing considerable

amounts of valuable data to be collected at that time. Once a carnivore is radio￾collared, data can be obtained about its movements, activity, home range or

territory, and dispersal. Often data about the creature’s predation and food habits

can also be collected, as well as information about its reproductive behavior. Several

chapters of this book deal with these subjects.

A subsidiary type of information, not directly related to a collared carnivore’s

movements, involves cause-specific mortality, including that from intraspecific

strife and diseases. Learning all this basic ecological, physiological, and behavioral

information then greatly aids in deriving mitigation measures for minimizing

depredation on livestock and other conflicts with humans, as well as facilitating

methods of restoring carnivores, monitoring the results, and furthering conserva￾tion efforts. Addressing those issues further rounds out this fine compendium.

Thus all in all, this book, edited by Luigi Boitani and Roger Powell, will be of

great use not only to carnivore researchers, but also to wildlife biologists through￾out the world who deal with carnivores, and it should stand as a milestone in the

carnivore-ecology and techniques literature for many years to come.

L. David Mech

US Geological Survey and University of Minnesota, USA

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vi | Foreword

Contents

List of contributors xv

1. Introduction: research and conservation of carnivores 1

Luigi Boitani and Roger A. Powell

2. Designing carnivore surveys 8

Luigi Boitani, Paolo Ciucci, and Alessio Mortelliti

2.1 Challenges of surveying carnivores 10

2.2 Planning a survey 10

2.2.1 Fundamentals of survey design: establishing goals and objectives 11

2.2.2 Fundamentals of survey design: carnivore survey data 12

2.2.3 Fundamentals of survey design: sampling design, methods,

and protocols 13

2.2.4 Fundamentals of survey design: statistically formalizing

survey objectives 14

2.3 Dealing with false absence 15

2.3.1 The fast growing family of occupancy models 16

2.3.2 Assumptions of occupancy models: the importance of a

priori planning 16

2.3.3 Some practical issues 18

2.3.4 Designing an occupancy study 18

2.4 Key issues for developing a survey design 19

2.4.1 Target population and spatial extent of the survey 19

2.4.2 Attribute to measure 20

2.4.3 Sampling design 21

2.4.4 Sampling effort 25

2.4.5 Tackling system variability: measures of precision and

their meaning 26

2.4.6 Field methods 29

3. Mind the map: trips and pitfalls in making and reading

maps of carnivore distribution 31

Carlo Rondinini and Luigi Boitani

3.1 Maps based on expert knowledge 32

3.1.1 Geographic range maps 32

3.1.2 Deductive habitat suitability models (HSM) 33

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3.2 Maps based on species’ occurrence surveys 34

3.2.1 Types of data 34

3.2.2 Biological significance and time relevance 36

3.2.3 Extrapolating points to map the distribution of a population 38

3.2.4 Inductive HSM 42

3.2.5 Caveats and limitations of deductive and inductive HSM 45

4. Noninvasive sampling for carnivores 47

Marcella J. Kelly, Julie Betsch, Claudia Wultsch, Bernardo Mesa, and L. Scott Mills

4.1 Methods of noninvasive sampling 48

4.1.1 Sign surveys 48

4.1.2 Genetic sampling 49

4.1.3 Camera-trap sampling 54

4.1.4 Endocrine/hormone sampling 55

4.2 Recent tools and advances in noninvasive sampling 56

4.2.1 Noninvasive DNA techniques 56

4.2.2 Using noninvasive DNA data 59

4.2.3 Data collection, handling, and analyses with remote cameras 62

4.2.4 Data collection, handling, and analyses for endocrine studies 65

4.3 Combining noninvasive and traditional approaches 67

4.3.1 Comparative approaches among noninvasive techniques 67

4.3.2 Combining traditional with noninvasive approaches 68

4.3.3 Data quality and integrity in noninvasive surveys 69

5. Humane and efficient capture and handling methods for carnivores 70

Gilbert Proulx, Marc R. L. Cattet, and Roger A. Powell

5.1 Mechanical capture methods 72

5.1.1 Traps and sets 72

5.1.2 Trapping efficiency 73

5.1.3 Humaneness 74

5.1.4 Traps and sets for specific carnivores 75

5.2 Use of drugs for capture and restraint of carnivores 78

5.2.1 Drug access, storage, and handling 78

5.2.2 Selection of drugs for use in carnivores 79

5.2.3 Methods to administer drugs 81

5.2.4 The value of knowledge and experience 81

5.3 Identification, prevention, and treatment of medical emergencies

associated with capture 84

5.3.1 Homeostasis, stress, distress, and treatment of medical emergencies 84

5.3.2 Necropsy 86

5.4 Euthanasia 86

5.5 Restraining and marking techniques 88

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viii | Contents

5.6 Designing effective trapping programs for carnivores 89

5.7 Animal welfare 89

Appendices 92

6. Carnivores in hand 130

Kerry R. Foresman

6.1 Aging 130

6.2 Standard body measurements 132

6.2.1 Body mass 133

6.2.2 Length measurements 133

6.2.3 Additional body measurements 136

6.2.4 Additional measurements, some to estimate age 137

6.2.5 Footpad patterns 137

6.3 Tooth eruption and measurements 137

6.3.1 Tooth eruption, wear, and age 138

6.3.2 Pulp cavity measurements and age 139

6.3.3 Cementum annuli and age 140

6.4 Skull and skeletal measurements 142

6.4.1 Skull measurements 142

6.4.2 Skull fusion and age 144

6.4.3 Skeletal morphology and age 144

6.4.4 Eye lens and age 144

6.5 Pelage and age 145

6.6 Sex and reproduction 145

6.7 Injuries 148

6.8 Physiological parameters 148

6.8.1 Blood 149

6.8.2 Tissue samples 149

6.8.3 Other samples 150

6.9 Bioelectrical impedance 150

6.10 Asymmetry 151

7. Radio-telemetry equipment and applications for carnivores 152

Mark R. Fuller and Todd K. Fuller

7.1 General background 152

7.2 Basic telemetry system 156

7.3 Radio-tracking field procedures 163

7.4 Satellite telemetry systems 163

7.5 Radio-telemetry applications for carnivores 166

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Contents | ix

8. Estimating demographic parameters 169

Ken H. Pollock, James D. Nichols, and K. Ullas Karanth

8.1 Combined challenges of carnivore ecology and survey logistics 170

8.2 Detection probabilities and demographic inference 171

8.3 Capture–recapture models 174

8.3.1 Closed models 175

8.3.2 Open models 177

8.3.3 Robust design models 178

8.3.4 Natural individual tags 178

8.3.5 Design of capture–recapture studies 180

8.4 Telemetry mortality models 180

8.4.1 Survival models 180

8.4.2 Combining telemetry and regular mark–recapture

models in one overall analysis 181

8.5 Occupancy models 181

8.5.1 Single-season models 181

8.5.2 Multi-season models 183

8.5.3 Software and study design 184

8.6 Probability sampling of carnivore tracks to estimate population density 185

8.7 Final thoughts 185

9. Movements, home ranges, activity, and dispersal 188

Roger A. Powell

9.1 Research design 189

9.2 Movements 191

9.3 Home range 193

9.4 Territories 196

9.5 Estimating animals’ home-ranges and territories 199

9.6 Home-range cores, overlap, and territoriality 208

9.6.1 Home-range cores 208

9.6.2 Home-range overlap 213

9.6.3 Static interactions 214

9.6.4 Dynamic interactions 215

9.6.5 Testing for territoriality 216

9.7 Parting thoughts 216

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10. Carnivore habitat ecology: integrating theory and application 218

Michael S. Mitchell and Mark Hebblewhite

10.1 What is habitat? 219

10.1.1 Potential, sink, quality, source, suitable, or critical?

What kind of habitat is it? 220

10.1.2 A fitness-based definition of habitat 222

10.2 What is carnivore habitat? 227

10.3 Measuring habitat use and selection by carnivores 232

10.3.1 The over-riding importance of questions 233

10.3.2 Why should carnivores be selective? 234

10.3.3 The importance of scale 236

10.3.4 Density dependence and habitat selection 237

10.3.5 Understanding habitat selection: study design 238

10.3.6 Using resource-selection functions and other approaches 240

10.3.7 Functional responses in resource selection 243

10.3.8 The importance of defining availability: recent advances

from the field of movement modeling 244

10.3.9 Quantifying resources 246

10.4 Linking habitat selection to population consequences 250

10.4.1 Habitat-based population estimates 251

10.4.2 Combining habitat and spatial models of mortality risk 252

10.4.3 Spatially explicit population models 253

10.5 Conclusions 255

11. Describing food habits and predation: field methods

and statistical considerations 256

Erlend B. Nilsen, David Christianson, Jean-Michel Gaillard, Duncan

Halley, John D.C. Linnell, Morten Odden, Manuela Panzacchi,

Carole Toı¨go, and Barbara Zimmermann

11.1 Quantifying predators’ diets 256

11.1.1 Scat analysis 256

11.1.2 Analysis of partly digested food items 259

11.1.3 Snow- and sandtracking 259

11.1.4 Telemetry-based methods to study predator diet 260

11.2 Ecological inferences from diet data 262

11.2.1 Quantifying kill rates and functional responses 262

11.2.2 Studying selection—the difference between use and availability 264

11.2.3 Quantifying food niche breadth and diet overlap 266

11.3 Using stable isotopes to infer trophic interactions 267

11.4 Estimating non-lethal effects of predation 269

11.5 Some further challenges 271

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Contents | xi

12. Reproductive biology and endocrine studies 273

Cheryl S. Asa

12.1 Carnivore reproductive physiology: the basics 273

12.1.1 Puberty 273

12.1.2 Seasonal reproduction 275

12.2 Stages of the female reproductive cycle 276

12.2.1 Pregnancy 278

12.2.2 Delayed implantation or embryonic diapause 279

12.2.3 Seasonal and lactational anovulation 280

12.2.4 Frequency of ovarian cycles 280

12.3 The endocrinology of stress 280

12.4 Endocrine studies and sampling strategies 281

12.5 Sample collection 284

12.5.1 Blood 284

12.5.2 Urine 285

12.5.3 Feces 286

12.5.4 Saliva 288

12.5.5 Hair 288

12.6 Non-endocrine techniques for studying reproduction 289

12.6.1 Males 289

12.6.2 Females 289

12.7 Gamete preservation and assisted reproduction 291

12.8 Control of reproduction 292

13. Investigating cause-specific mortality and diseases in

carnivores: tools and techniques 294

Greta M. Wengert, Mourad W. Gabriel, and Deana L. Clifford

13.1 Determining causes of mortality in carnivores 294

13.1.1 Locating dead animals to determine cause-specific mortality 295

13.1.2 Handling dead animals and important precautions 296

13.1.3 Field-data collection at mortality sites 296

13.1.4 The clinical necropsy 297

13.1.5 When clinical necropsies just aren’t feasible—a

quick guide to field necropsy 298

13.1.6 Field and laboratory investigation of intraguild predation 298

13.2 Studying disease and pathogen cycles in carnivores 300

13.2.1 Detection of disease, infection, and pathogen exposure 300

13.2.2 Epizootiology in carnivore populations 305

13.2.3 Modeling techniques in disease ecology 308

13.3 Prevention and control of disease 310

13.3.1 Intervention options: removing the causative factor 310

13.3.2 Intervention options: manipulating the host population 310

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xii | Contents

13.3.3 Intervention options: manipulating sympatric species

including domestic animals 312

13.3.4 Intervention options: addressing human activities 313

14. Mitigation methods for conflicts associated with carnivore

depredation on livestock 314

John D. C. Linnell, John Odden, and Annette Mertens

14.1 Who kills whom? 315

14.2 Documenting depredation 315

14.3 The ecology of depredation and its mitigation 319

14.3.1 Avoiding encounters between carnivores and livestock 319

14.3.2 Preventing the recognition of livestock as potential prey 321

14.3.3 Preventing access to livestock by carnivores 324

14.4 Compensation 328

14.5 Integrating mitigation into agricultural policy 330

15. Carnivore restoration 333

Michael K. Stoskopf

15.1 Human dimension 334

15.1.1 Cultural issues 334

15.1.2 Political and jurisdictional issues 335

15.1.3 Economics 335

15.2 Environmental and habitat dimension 337

15.2.1 Topography 337

15.2.2 Climate 338

15.2.3 Anthropogenic features 339

15.2.4 Prey base 339

15.2.5 Health-risk assessment 339

15.3 Animal dimension 340

15.3.1 Carnivore–carnivore interactions 340

15.3.2 Carnivore-prey interactions 342

15.3.3 Selecting founder populations 342

15.3.4 Use of captive animals for restoration 342

15.3.5 Genetic management 344

15.3.6 Hybridization and introgression management 345

15.3.7 Health management and biosecurity 346

15.3.8 Health interventions 347

15.3.9 Adaptive management 348

15.3.10 Release methods 349

15.3.11 Population augmentation 350

15.4 Exit strategy 351

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Contents | xiii

16. Designing a monitoring plan 353

Eric M. Gese, Hilary S. Cooley, and Frederick F. Knowlton

16.1 Identifying questions and monitoring designs 354

16.2 Developing a monitoring program 355

16.3 Evaluating the monitoring plan 358

16.3.1 Thresholds and trigger points 358

16.3.2 Forecasting trends 359

16.3.3 Predicting patterns over space and time 359

16.3.4 Integrating monitoring data 360

16.3.5 Risk analysis 360

16.4 Changing the monitoring plan 360

17. Assessing conservation status and units for conservation 362

Urs Breitenmoser, Christine Breitenmoser-Wu¨rsten, and Luigi Boitani

17.1 Assessing extinction risks for carnivore populations 363

17.2 Identifying and delineating carnivore conservation units 368

17.2.1 Choosing biological entities 368

17.2.2 Socio-political considerations 371

17.2.3 Geographic delineation 373

17.3 Designating and establishing carnivore conservation units 375

17.4 Final thoughts 378

References 379

Index 491

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xiv | Contents