K.
Ullas Karanth is a Conservation
T he tiger has served as an effective
flagship species in the global efforts
to conserve a variety of wildlife and
wild-lands in Asia. However, despite
the high public concern for tigers
and the consequent conservation
investments during the past three
decades, wild tigers are still gravely
threatened. Their preservation
requires, more than ever before,
conservation interventions guided
by reliable ecological knowledge.
Such knowledge can only be
generated by the application of
rigorous methods of data collection.
This manual presents a conceptually
unified, statistically rigorous
approach to ecological monitoring
of wild tigers. It synthesizes current
scientific understanding of tiger
ecology with recent advances in
population sampling techniques. The
authors of this manual present an
array of approaches for monitoring
populations of tigers and their
principal prey species: these methods
are tailored to suit a variety of
practical contexts. The topics
covered in this illustrated manual
include methods for the rigorous
estimation of habitat occupancy as
well as estimation of relative and
absolute densities. This manual will
be a useful reference for wildlife
researchers, managers and
conservationists.
Indian Rupees 400/-
Printed in India
Scientist with Wildlife Conservation
Society (WCS), New York and is the
Director of its India Program. He
received his undergraduate degree in A Manual for
engineering from Mysore University, Wildlife Researchers,
India, his Masters degree in wildlife
biology from the University of Florida, USA, and his Managers and
Doctorate in applied zoology from Mangalore University, Conservationists in
India. He has focused his research on the ecology of tigers, Tropical Asia
other carnivores and their prey species. He is especially
interested in developing monitoring techniques and in Edited by K. Ullas Karanth and James D. Nichols
conservation policy issues. Ullas Karanth has published
over 50 scientific articles and is a fellow of the Zoological
Society of London. He has been internationally recognized
for his contributions to tiger conservation and has served
on the steering committee of the Indian Government’s Project
Tiger.
James D. Nichols is a senior scientist
working with the U.S. Geological
Survey’s Patuxent Wildlife Research
Center. He received his undergraduate
degree in biology from Wake Forest
Edited by K. Ullas Karanth and James D. Nichols
University, Masters degree in wildlife
management from Louisiana State
University and his Doctorate in Wildlife Ecology from
Michigan State University, in USA. His research interest
has been in animal population dynamics and wildlife
management with a special focus on estimation of
demographic parameters. He has published over 200
scientific articles and is widely recognized for his theoretical
contributions in this field. James Nichols is the recipient of
various awards including the Outstanding Publication Award
and the Outstanding Monograph Award from The Wildlife
Society, the George W. Snedecor Award from the American
Statistical Association, and the best research paper award
from US Fish and Wildlife Service.
Centre
for
Wildlife
Studies
 MONITORING TIGERS AND THEIR PREY
A Manual for Wildlife Researchers, Managers and Conservationists in Tropical Asia

Edited by

K. ULLAS KARANTH and JAMES D. NICHOLS

2ND COVER

Centre for Wildlife Studies

Published by Centre for Wildlife Studies, CONTENTS
26-2, Aga Abbas Ali Road (Apt: 403), Bangalore, Karnataka-560 042, India
Email: wcsind@bgl.vsnl.net.in
Foreword By John Seidensticker ix
© Centre for Wildlife Studies 2002
Acknowledgements xii
This book is in copyright. Subject to statutory exception and to the provisions of the relevant
collective licensing agreements, no reproduction of any part may take place without the written Preface xiv
permission of Centre for Wildlife Studies.
CHAPTER 1
First published in India 2002 Monitoring Tigers and Prey: Conservation Needs and
ISBN 81-901442-1-9 Managerial Constraints 1

Title: Monitoring Tigers and Their Prey: K. Ullas Karanth, James D. Nichols, P. K. Sen and Vinod Rishi
A Manual for Researchers, Managers and Conservationists in Tropical Asia - Introduction
- The sampling-based approach to monitoring
Editors: K. Ullas Karanth and James D. Nichols - Defining objectives
- Assessing available resources
Picture Credits (Cover): K. Ullas Karanth and Praveen Bhargav - Decision making: Matching objectives and resources
- References
Printed in India.
CHAPTER 2
All brand and product names referred in this manual are registered trade marks of their respective companies.
Ecology of the Tiger: Implications for Population Monitoring 9

K. Ullas Karanth and Raghunandan S. Chundawat

Support for the publication of this book has been provided by the following organizations
- Introduction
- Tiger social organization and land tenure system
- Ecological determinants of tiger density
Wildlife Conservation Society - Survival, mortality and population dynamics
International Programs - Effect of environmental factors on monitoring
2300, Southern Boulevard, Bronx, New York, NY-10460-1099, USA. - Tiger ecology: Challenges to monitoring
www.wcs.org - How to monitor tigers
- The need for a unified framework for monitoring
- References

US Geological Survey CHAPTER 3

Patuxent Wildlife Research Center, Laurel, MD-20708-4017, USA.
www.pwrc.usgs.gov Population Monitoring: A Conceptual Framework 23

James D. Nichols and K. Ullas Karanth

- Introduction
World Wildlife Fund-US - The statistical framework
1250, 24th Street NW, Washington DC-20037, USA. - References
www.worldwildlife.org

iii
 Contents Contents

CHAPTER 4 CHAPTER 8
Statistical Concepts: Assessing Spatial Distributions 29 Field Surveys: Assessing Relative Abundances of Tigers and Prey 71

James D. Nichols and K. Ullas Karanth K. Ullas Karanth and N. Samba Kumar

- Presence-absence surveys - Index surveys

- Mapping the spatial distribution - Indices of relative abundance for tigers
- Looking ahead - Indices of relative abundance for prey species
- References - References

CHAPTER 5 CHAPTER 9
Field Surveys: Assessing Spatial Distributions of Tigers and Prey 39 Statistical Concepts: Estimating Absolute Densities of Prey
Species Using Line Transect Sampling 87
K. Ullas Karanth, N. Samba Kumar and Raghunandan S. Chundawat
Len Thomas and K. Ullas Karanth
- Introduction
- Field surveys - Introduction
- Questionnaire surveys - Line transect theory
- Organizing the survey data for mapping and analysis - Assumptions of line transect sampling
- References - Survey design
- Data analysis
CHAPTER 6 - References

Spatial Distributions of Tigers and Prey: Mapping and CHAPTER 10

the Use of GIS
Bradley M. Stith and N. Samba Kumar
- Introduction
- Data sources
- Compiling survey data in the GIS
- Ancillary sources of GIS data
- Summarizing data and querying the GIS database
- Producing maps of spatial distribution
- Analyzing tiger-prey-environment relationships
- References
51
Field Surveys: Estimating Absolute Densities of Prey Species
Using Line Transect Sampling 111
K. Ullas Karanth, Len Thomas and N. Samba Kumar
- Introduction
- What are “True” transect lines?
- Placement of transects: Practical considerations
- Meeting line transect assumptions in field surveys
- Season and timing of line transect surveys
- References

CHAPTER 7 CHAPTER 11
Statistical Concepts: Indices of Relative Abundance 61 Statistical Concepts: Estimating Absolute Densities of Tigers
Using Capture-Recapture Sampling 121
James D. Nichols and K. Ullas Karanth
James D. Nichols and K. Ullas Karanth
- Kinds of indices
- Reducing bias in index-based estimates - Introduction
- Robust estimation of relative abundance - Photo trapping tigers
- Summary comments on indices and estimation - Capture-recapture models
- References - Survey design considerations

iv v
 Contents Contents

- Conclusion APPENDIX 3.0

- References
Specimen Data Form for Line Transect Surveys 182
CHAPTER 12
APPENDIX 4.0
Field Surveys: Estimating Absolute Densities of Tigers Using
Capture-Recapture Sampling 139 Specimen Data Form for Camera Trap Survey of Tigers 183

K. Ullas Karanth, N. Samba Kumar and James D. Nichols APPENDIX 5.0

- Introduction Design of the Javaji Metal Secure Camera Trap Shell 184
- Choice of camera trap equipment
- Conducting camera trap surveys APPENDIX 6.0
- Analysis of capture-recapture data
- References Web Sites for Free Analytical Software and Addresses of some
Camera Trap Equipment Suppliers 187
CHAPTER 13
APPENDIX 7.0
Monitoring Tiger Populations: Why Use Capture-Recapture
Sampling? 153 Glossary of Technical Terms 189

K. Ullas Karanth and James D. Nichols APPENDIX 8.0

- The natural advantage Common and Scientific Names of Animals Used in the Text 193
- References

APPENDIX 1.1
Drawings of the Tiger and other Sympatric Large
Carnivores and their Tracks 167

APPENDIX 1.2
Drawings of Principal Prey Species of the Tiger and their Tracks 170

APPENDIX 1.3
Specimen Data Form for Field Surveys of Spatial Distribution of
Tigers and Prey 176

APPENDIX 1.4
Specimen Data Form for Questionnaire Surveys of Informants 178

APPENDIX 2.1
Specimen Data Form for Encounter Rate Surveys of Tiger Sign 180

APPENDIX 2.2
Specimen Data Form for Recording Dung Counts on Plots 181

vi vii
 FOREWORD

“Assessment of change, dynamics, and cause and effect are at the heart of thinking and
explanation. To understand is to know what causes provoke what effects by what means, at
what rate. How then is such knowledge to be represented?”

Edward Tufte
[Tufte, E. D. 1997. Visual Explanations: Images and quantities, evidence and narrative. Graphics press, Cheshire, CN, USA.]

Tigers are breathtaking, marvelous, and provoke strong feelings where and whenever you are
privileged to see one. Tigers are the specialized predators of large ungulates. Because tigers
are secretive, prefer thick cover, and live at large landscape scales, counting them with
confidence is challenging and few scientists have managed to do so. But do so we must if we

BLANK are to advance our understanding of tiger ecology, predict how stresses and changes in the
ecosystems where they live impact them, and explore how tigers, as the top carnivore in
Asian forests, themselves control ecosystem processes.

I recently received an urgent request from a teacher of our next generations of wildlife scientists
at the Wildlife Institute of India, Ravi Chellam. He asked the following: “1) How many
habitat fragments with confirmed connectivity for genetic exchange in India support tigers?
Can TCUs (Tiger Conservation Units) be taken as areas with an intermingling tiger population?
How many TCUs do we have in the country? 2) How many protected areas and of what
status are in these fragments/TCUs? 3) What is the average size, and what is the largest size
of tiger population numbers in the various fragments/TCUs? 4) How many tiger populations
with confirmed genetic exchange will number more than 200, more than100, and less than
50? 5) What is your estimate for the number of tigers outside PAs? 6) What is your best
estimate for India’s tiger population?”

Good questions. It is essential that we are able to answer these and many more questions if we
are to understand and overcome the barriers to securing a future for wild tigers. But even
after more than 30 years of tiger conservation efforts in India by many people who have
devoted their lives to securing a future for wild tigers, we cannot answer these questions in
detail. I could only refer Ravi Chellam to the best source of information I have in hand: A
Framework for Identifying High Priority Areas and Actions for the Conservation of Tigers
in the Wild, a report jointly prepared by Wildlife Conservation Society and World Wildlife

ix
 Foreword Foreword

Fund, USA in1997. The situation in most other tiger range nations is about the same.
Recently, however, we are seeing progress in our ways of knowing how tigers respond to
changing conditions. For example, Dale Miquelle and his associates have advanced a sampling-
based, quantitative monitoring program for tigers in the Russian Far East. Timothy O’Brien
and colleagues have proposed one for rainforests in Indonesia. Now, Ullas Karanth, James
Nichols, and their co-authors have provided wildlife scientists and managers with this
comprehensive account of how to scientifically monitor tigers and their prey in tropical Asian
landscapes.
One of our deepest historical traditions is the assumption that knowledge comes in the form
of a hierarchy. The idea that knowledge is a hierarchy has persisted for 2,500 years but the
form of knowledge privileged to perch at the very top has changed over time. The nature of
scientific inquiry has changed as our ability and the tools that we use to measure and quantify
have advanced. Most of us in the Western tradition of science work with the assumption that
knowledge is ordered in this hierarchical fashion with science at the top, professional and
scholarly or “expert” knowledge in the middle, and the opinions of the mass public at the
bottom as noted by eminent biologist Ernst Mayr.
best statistical applications: Nor did they have transparency in their data collection and analysis.
This monitoring work could not be replicated. It was not subject to peer review. It simply
wasn’t science.
Editors Ullas Karanth and James Nichols and their co-authors explain in the pages that
follow, how the technology and methodology for counting tigers and their prey in non-invasive
ways have greatly advanced in the last two decades, to the point that monitoring tiger and
prey numbers can escape the realm of expert opinion and become science-based. We can now
begin to institute a program to answer the critical questions posed by my colleague from the
Wildlife Institute of India. All of this is essential knowledge that enables and enhances our
efforts to secure a future for wild tigers.
JOHN SEIDENSTICKER
Chairman, Save The Tiger Fund Council
Senior Curator, Smithsonian National Zoological Park

We have seen the statements, over and over, in the general press that at the turn of the century
there were 100,000 tigers in the wild in Asia and there were 40,000 tigers in India. There was
nothing scientific in the way these numbers were derived. They were given their credibility
and they have been retained because they were based on more than mass public knowledge;
these numbers were “expert” opinion, the second rung in the knowledge hierarchy. At the
beginning of Project Tiger, its director, K.S. Sankhala derived an estimate of tiger numbers
based on his experience. J.C. Daniel of the Bombay Natural History Society used a
questionnaire survey of forest officers to estimate tiger numbers in India. Both came to similar
conclusions (about 2,000 tigers), and were given credibility because they too were based on
expert opinion. Deriving these estimates had nothing to do with science.

Counting tigers in India using ‘pugmark censuses’ was an effort to bring science into the
picture and to replace expert opinion. ‘Pugmark census’ was meant to be objective, and take
counting tigers to the next higher rung, to the apex of our hierarchy of knowledge. The
problem was that the proponents of counting tigers via ‘pugmark census’ did not follow the
usual conventions of scientific inquiry and include optimum sampling procedures with the

x xi
 Acknowledgements

ACKNOWLEDGEMENTS
While we worked on this manual, Wildlife Conservation Society, New York, supported Ullas
Karanth and US Geological Survey supported James Nichols. Subsequently, World Wildlife
Fund-USA, Washington DC and US Geological Survey, Patuxent Wildlife Research Center,
Laurel, MD, USA provided support for its publication. We are extremely grateful to the
above three institutions.
Bengal, Assam and Arunachal Pradesh. Besides administratively facilitating our research
projects, Centre for Wildlife Studies, Bangalore, India, also published this manual. We are
very grateful to all the above institutions and their individual staff members who supported
our work.
K. ULLAS KARANTH JAMES D. NICHOLS

This book has benefited from intellectual and material support we received from several
individuals and institutions. As editors, it is our duty to acknowledge all these contributions.
We are indebted to John Seidensticker who wrote the foreword, and, the co-authors who
contributed to different chapters: Samba Kumar; Len Thomas; Raghu Chundawat; Brad
Stith; P. K. Sen and Vinod Rishi. We acknowledge Maya Ramaswamy who prepared the
illustrations; Abi Tamim who assisted in the final editing; Praveen Bhargav and R. S. Suresh
who designed the layout of the book and Bonnet Designs for printing and production.

Over the past several years, and, in the course of developing the contents of this manual, we
have benefited enormously from the unstinting assistance we received - in the field and off it
- from K. M. Chinnappa, Samba Kumar and Jim Hines, to whom we are especially indebted.

The following participants at the Wildlife Conservation Society sponsored Nagarahole tiger
monitoring workshop in 1999 provided useful comments on the ideas presented here: Alexander
Hiby; Anil Gore; Sharayu Paranjpe; Tim O’Brien; Kae Kawanishi; M. B. Krishna;
M. D. Madhusudan; Ravi Chellam; Yadavendradev Jhala; Qamar Qureshi; Valmik Thapar;
A. Udhayan; Vijay Kumar Gogi and N. Rajasekhar. We thank all these colleagues and the
various institutions that sponsored their participation in the Nagarahole workshop.

Over the years, the following institutions have provided sustained support for our research
projects on tiger ecology. In the United States of America, these institutions include : Wildlife
Conservation Society; US Fish and Wildlife Service (Division of International Conservation);
US Geological Survey (Patuxent Wildlife Research Center), and Save the Tiger Fund of the
National Fish and Wildlife Foundation and ExxonMobil Corporation. In the United Kingdom,
support was provided by Global Tiger Patrol. In India, our work has been facilitated by: The
Ministry of Environment and Forests (Directorate of Project Tiger and the US-India Fund);
The State Forest Departments of Karnataka, Madhya Pradesh, Maharashtra, Rajasthan, West

xii xiii
 Preface

PREFACE We strongly believe that the primary yardstick of the reliability of any animal monitoring
methodology should be the process of scientific peer review and publication. Tiger biologists,
The tiger is an endangered animal that has served as an effective flagship species in conserving managers and conservationists cannot enter special pleadings to claim exemption from this
wildlife and wild-lands in many parts of Asia. For over a quarter century, there has been a universal scientific norm. Therefore, we have chosen to present in this book only those
high level of public concern for tigers, leading to major investments in conservation efforts. monitoring approaches that are adequately justified by peer-reviewed scientific literature: We
However, wild tigers continue to be under grave threat, and their preservation now requires, make no apologies for this. However, we are well aware that, eventually, the methods that we
more than ever before, informed conservation interventions guided by reliable ecological present in this manual will be replaced by better methods generated through the very same
knowledge. Such reliable knowledge can only be generated through application of rigorous process of scientific review that we endorse. In fact, we look forward to such improvements.
methods of data collection. The lack of scientific rigor in the approaches to assess the status
of wild tigers and prey is now clearly recognized as a serious gap in global conservation In the meanwhile, we hope that the methods we present in this manual will satisfy the current
efforts. Therefore, a comprehensive technical manual that describes methods useful for needs of its targeted audience, both in terms of scientific rigor and practical utility.
monitoring populations of wild tigers and their prey species is a major conservation need at
the moment.
K. ULLAS KARANTH JAMES D. NICHOLS
At our initiative, Wildlife Conservation Society (WCS) tried to address this need by sponsoring
an international technical workshop in Nagarahole Reserve, India, during January 1999.
This workshop on techniques for monitoring tiger and prey populations was conducted in
association with the Directorate of Project Tiger of the Indian government as well as state
forestry departments. The participants in the workshop included experienced wildlife managers,
carnivore biologists and bio-statisticians specializing in animal population estimation issues.
This manual is based on some ideas presented at the workshop by the contributing authors.
Its goal is to present a conceptually unified, rigorous approach to monitoring tiger and prey
populations, synthesizing current scientific understanding of tiger ecology with recent advances
in population estimation methods. At the same time, the authors have tried to present the
reader with an array of monitoring approaches that are tailored to a variety of practical
contexts.

In practical terms, tiger habitats in Asia fall under two major categories: The temperate zone
forests in the Russian Far East and China, and the tropical forests in South and Southeast
Asia. In the Russian Far east, the presence of a uniform substrate of snow, extremely low
tiger densities, and, the extensive geographic scale of monitoring set the technical issues
clearly apart from those in tropical Asia. We have chosen to restrict the scope of this manual
to primarily deal with tiger monitoring issues in tropical Asia that we are familiar with.
However, some of the basic concepts and techniques outlined here are likely to be useful for
monitoring tigers in temperate zone forests also.

xiv xv
 CHAPTER 1

MONITORING TIGERS AND PREY: CONSERVATION NEEDS

AND MANAGERIAL CONSTRAINTS

K. Ullas Karanth1, James D. Nichols2, P. K. Sen3 and Vinod Rishi4

Wildlife Conservation Society (International Programs), Bronx, New York, NY-10460-1099, USA.
1

Email: <ukaranth@wcs.org>

United States Geological Survey, Patuxent Wildlife Research Center, Laurel, MD-20708-4017, USA.
2

Email: <jim_nichols@usgs.gov>

World Wide Fund for Nature-India, 172-B Lodi Estate, New Delhi-110 003, India.
3

Email: <wwftcp@vsnl.com>

Indira Gandhi National Forest Academy, P.O. New Forest, Dehra Dun, Uttaranchal-248 006, India.
4

Email: <director@ignfa.up.nic.in>

BLANK INTRODUCTION

The tiger has been used as a flagship species of wildlife conservation efforts in several Asian
countries since the early 1970’s. For example, India’s “Project Tiger”, initiated in 1973 is
recognized as one of the largest and most effective endangered species recovery projects
undertaken in developing countries. Since then, Asian countries have invested millions of
dollars in tiger conservation efforts. However, given the massive pressures imposed on tigers,
through over-hunting of their prey species, poaching of tigers and damage to habitats from a
variety of causes (Karanth 2001), wild tiger populations are not yet on a secure footing. It is
likely that tigers will be under threat for many years to come even as conservation efforts
continue.

The need for scientific monitoring of tiger and prey populations arises from three considerations.
Firstly, there is the need to objectively evaluate the success or failure of management
interventions, so as to react adaptively and solve problems. Clearly this is the primary goal of
wildlife managers and conservation agencies. A second major goal of scientific monitoring of
tiger and prey populations is to establish benchmark data that can serve as a basis for future
management. Our management goals and expectations are often heavily influenced by our
perceptions of pre-management population status. Such development of scientific monitoring
is primarily the task of well-trained wildlife biologists or managers who are seriously interested
in collecting benchmark data. A third broader goal of tiger monitoring is to develop a body of
empirical and theoretical knowledge that can potentially improve our predictive capacity to
deal with new situations. Such basic science also contributes to the general advancement of

1
 Monitoring Tigers and Prey: Conservation Needs and Managerial Constraints
human knowledge. This task falls largely in the domain of academic studies carried out by
wildlife biologists.
Given the tiger’s critical status, and, the substantial investments being made in tiger
conservation, wildlife managers and conservation agencies need clear and reliable answers to
some basic questions. Without these answers, they cannot even begin to evaluate the success
or failure of conservation efforts. Some of these basic questions are:
1. What is the distributional range occupied by individual tiger populations?
2. Where are individual tiger populations increasing their range, and, where are these ranges
fragmenting or shrinking?
3. Within the widespread distributional range of tigers, what is the proportion of the area
occupied by productive, breeding populations?
4. In important individual tiger reserves, what are the tiger population trends? Are tiger
populations in such reserves holding steady, declining or increasing?
The traditional responses to such questions have been based on trying to obtain total counts
of wild tigers through ‘censuses’ (Choudhury 1970, 1972). Such ‘census-based’ approaches
have many biological and statistical weaknesses (Karanth 1987, 1999; Karanth et al. in
prep.). Consequently, it is now officially recognized that more reliable, ‘population sampling-
based’ methods are needed to monitor tiger populations.
During the past 35 years, our knowledge of tiger ecology has advanced significantly as a
result of several scientific studies (Chapter 2). Some of these studies have applied
advanced tools such as radio telemetry and camera trapping to gain new information and
insights.
During the past three decades, concepts and methods used for assessing wildlife population
parameters have also made great progress (Chapter 3). Two important conceptual approaches
to population sampling - Distance Sampling and Capture-Recapture Sampling, have advanced
particularly rapidly (Buckland et al. 1993, 2001; Nichols 1992; Thompson et al. 1998).
They now offer powerful tools for monitoring wild animal populations.
As a result, it is now recognized that methods we use for monitoring tigers and prey species
must also incorporate recent scientific advances. As an example of such recognition, India’s
Project Tiger Directorate recently issued new guidelines that attempt to revise the traditional
monitoring protocols (Anon. 2001).
2 3
Monitoring Tigers and Prey: Conservation Needs and Managerial Constraints
Over the years, the editors of this manual, together with other collaborators, have developed
several sampling-based approaches to estimating population parameters of tigers and their
prey species (Karanth and Sunquist 1992; Karanth 1995; Karanth and Nichols 1998, 2000).
In this context, the authors believe that a comprehensive manual incorporating these new
approaches would be very useful for tiger biologists, reserve managers, conservation agencies
and individual conservationists engaged in monitoring wild tigers.
THE SAMPLING-BASED APPROACH TO MONITORING
We recognize the need to prevent this manual from turning into an exercise in ‘reinventing the
wheel’ by trying to review all established scientific methods of animal population estimation:
There are several excellent general manuals that deal with these issues (Seber 1982; Bookhout
1994; Rabinowitz 1997; Thompson et al. 1998; Williams et al. 2002). We expect the users of
this manual to familiarize themselves with basic statistical methods and standard literature
on population estimation. However, for the sake of completeness, brief overviews of current
sampling-based population estimation methods are provided in this manual.
More importantly, the seemingly disparate approaches to counting animals are unified (Chapter
3), under a rigorous conceptual framework established in the standard works of reference
(Seber 1982; Thompson 1992; Lancia et al 1994; Thompson et al. 1998; Williams et al.
2002). Readers are directed to these sources for details.
Specifically, this manual will consider the manner in which monitoring approaches deal with
the core problems of spatial sampling and observability. Spatial sampling concerns the frequent
inability to use animal survey methods over an entire area of interest. In such cases, we need
to survey some subset of the entire area of interest and then use these results to draw inference
about the entire area. Observability concerns the typical inability to detect and count all
animals present in an area that is selected for survey.
Regardless of the particular survey method, comparisons of resulting count statistics over
time or space require consideration of the associated detection probabilities (probability that
an animal appears in the count statistic). Some approaches to animal monitoring permit
direct estimation of these detection probabilities (these methods tend to be those with the
greatest requirements for effort and resources) and permit strong inferences, whereas others
rely on strong assumptions about the absence of variation in these probabilities over time
and/or space and typically yield weaker inferences. Focus on these key features of animal
monitoring will facilitate useful consideration and comparison of the various approaches
suggested for monitoring tigers and their prey.
 Monitoring Tigers and Prey: Conservation Needs and Managerial Constraints
DEFINING OBJECTIVES
The very first step in monitoring any tiger/prey population is to define the objectives of the
exercise that you want to undertake. These specific objectives are linked to the three broad
monitoring goals we identified at the beginning of this chapter. Any monitoring program can
have one or more of the following specific objectives:
1. Mapping the distribution of tigers and principal prey species on a regional and countrywide
basis.
2. Estimating population trends in order to understand whether populations are increasing or
decreasing in selected reserves. Such surveys frequently use quantitative indices of relative
abundances of tigers and prey species (e.g., number of tiger track sets seen per 100 km
walked by the observer; number of sambar pellets found in a 100 m2 plot).
3. Estimating the absolute densities of tigers and prey (e.g., number of tigers/100 km 2 or
number of sambar/1 km2) in high priority protected areas.
4. Estimating the rates of annual survival, recruitment and population change through long
term studies.
ASSESSING AVAILABLE RESOURCES
Which of the objectives outlined above can be achieved depends on the time, manpower,
technical skills and material resources that are at your command. Therefore, assessing the
resources available to you is the second important step of tiger monitoring. Usually monitoring
of tigers and prey is carried out either by government agencies (e.g., forestry or wildlife
departments) or by wildlife biologists conducting focused research or surveys. The survey
personnel available may vary greatly in terms of their technical skills and field abilities. Their
numbers may range from a handful of highly trained naturalists to dozens or even hundreds
of field personnel without scientific skills.
The skills required for carrying out field surveys of tigers and prey also vary greatly. “Field
skills” like the ability to walk long distances, observe carefully, recognize and record animals
or their signs accurately, are of prime importance. Persons with such field skills may be
wildlife biologists, wildlife department staff or local hunters/naturalists. Very different kinds
of skills are necessary for designing the surveys and for analyzing the data that result from
the field surveys. We will call these “analytical skills”. These skills include knowledge of
population sampling methods, ability to organize and analyze the field survey data and to
4 5
Monitoring Tigers and Prey: Conservation Needs and Managerial Constraints
interpret the results correctly. It is critically important for you to assess the kinds of skills that
are available (or absent) in your particular situation.
Similarly, the material resources available for tiger monitoring vary depending on the local
context. In most cases, particularly where wildlife or forestry departments carry out the tiger
monitoring, only basic tools may be available. Such “basic tools” may include: Rudimentary
maps of the area being surveyed, plaster of Paris (Calcium sulphate) powder, glass tracers,
plastic ropes, polythene bags, identification tags etc. These “basic tools” are absolutely essential
to carry out any monitoring program. Sometimes, in addition to basic tools, “special tools”
such as compasses, cameras, range finders and global positioning systems (GPS) may be
available. Rarely, even “advanced tools” such as camera traps, radio telemetry equipment,
computers and special software may also be available.
DECISION-MAKING: MATCHING OBJECTIVES AND RESOURCES
A successful monitoring program depends on your being able to come up with a survey
scheme that defines the achievable objectives carefully, based on available manpower, technical
skills and material resources. Setting up wonderful goals based on mere wishful thinking is
not useful. The goals must be realistic in your specific context; if not, the failure of your
monitoring program is almost guaranteed.
You should examine the resources available at each administrative unit/level (State, Region,
Reserve). The type of monitoring feasible can be determined by matching the available
resources, local ecological conditions and potential survey methods. The following guidelines
may be useful to select the survey methods that you can employ reliably:
1. Recognize that it is simply impossible to achieve some of the objectives defined earlier
(relative or absolute density estimation for tiger and prey species) if resources are extremely
limited. Such limitation will nearly always occur when dealing with large regions, states
or countries. Therefore, if you have access to only basic tools and untrained field personnel,
you can only do surveys of presence of tigers and prey species (Objective 1), using the
concepts and methods described in Chapters 4, 5 and 6. This is likely to be the situation
that you will encounter over 90% of the tiger distributional range in tropical Asia.
2. If you have trained wildlife biologists or personnel, you can try to get relative density
estimates of tigers from track or scat-based indices and relative density estimates of prey
species using pellet or dung counts (Objective 2), in addition to meeting Objective 1. It is
likely that you can do such index-based surveys only in individual reserves or study
 Monitoring Tigers and Prey: Conservation Needs and Managerial Constraints
areas. The concepts and methods used for attaining these are described in Chapters 7
and 8.
3. If you have access to some special tools, enough trained personnel and wildlife biologists,
then, in addition to Objectives 1 and 2, you can estimate absolute densities of
prey species (Objective 3), using the Distance-sampling concepts and methods detailed in
Chapters 9 and 10. Such surveys are also likely to be feasible only in individual reserves
or study areas.
4. If you have access to all the above resources, plus an adequate number of camera traps
(advanced tools), then you can do camera trap capture-recapture sample surveys of tigers
using methods described in Chapters 11 and 12, to estimate absolute densities. This type
of survey may be feasible only in a few selected study areas.
However, it is likely that over 90% of the tiger distributional range, the sample sizes attained
from field counts of tiger tracks, camera trap photos and prey dung, etc., may be too low for
deriving density estimates or even precise indices. Therefore, you may have to be content
with focusing only on Objective (1) above: Mapping the distribution of tigers and principal
prey over much of tiger range.
If there is a serious mismatch between resources available and objectives you hope to attain,
please do not attempt to meet goals 2 or 3. There is absolutely nothing wrong with this:
Monitoring of tiger distribution is a critically important first step in implementing any landscape
level conservation program. In such a case we recommend that you initiate your survey
efforts with goal 1. Gradually, over the years, you can build up the capacity and resources to
try to meet Objectives 2 and 3.
We emphasize that if you really want to estimate parameters such as population size,
absolute densities, survival and recruitment rates, you have to employ advanced methods
described in Chapters 9 to 12. However, we note that these are primarily expensive
research tools and can be employed only in reserves where skilled manpower and
resources are available. Such advanced techniques cannot be applied for routine population
surveys over the entire tiger distribution range. The advanced methods are relevant for
monitoring tigers over a very small fraction of the animal’s vast range. On the other hand, the
two most critical needs of tiger and prey monitoring in Asia, mapping spatial distributions
over large regions and determining population trends in specific reserves through
indices, are widely attainable goals using the relatively simple methods outlined in
Chapters 4 to 8.
6 7
Monitoring Tigers and Prey: Conservation Needs and Managerial Constraints
REFERENCES
Anonymous, 2001. Guidelines for monitoring tiger and prey populations. Project Tiger
Directorate, Ministry of Environment and Forests, Government of India, New Delhi
(Unpublished).
Bookhout, T. (Ed.). 1994. Research and management techniques for wildlife and habitats.
The Wildlife Society, Bethesda, MD, USA.
Buckland, S.T., Anderson, D.R., Burnham, K.P. and Laake, J.L. 1993. Distance sampling:
Estimating abundance of biological populations. Chapman and Hall, London, UK.
Buckland, S.T., Anderson, D.R., Burnham, K.P., Laake, J.L., Borchers, D.L. and Thomas,
L. 2001. Introduction to distance sampling. Oxford University Press, Oxford, UK.
Choudhury, S.R. 1970. Let us count our tigers. Cheetal 14:41-51.
Choudhury, S.R. 1972. Tiger census in India. Cheetal 15:67-84.
Karanth, K.U. 1987. Tigers in India: A critical review of field censuses. Pages 118-133 in
Tigers of the world: The biology, biopolitics, management and conservation of an
endangered species (Eds: R.L. Tilson and U.S. Seal). Noyes Publications, Park
Ridge, NJ, USA.
Karanth, K.U. 1995. Estimating tiger populations from camera-trap data using capture-
recapture models. Biological Conservation 71:333-338.
Karanth, K.U. 1999. Counting tigers, with confidence. Pages 350-353 in Riding the tiger:
Tiger conservation in human-dominated landscapes (Eds: J. Seidensticker, S. Christie
and P. Jackson). Cambridge University Press, Cambridge, UK.
Karanth, K.U. 2001. The way of the tiger: Natural history and conservation of the endangered
big cat. Voyageur Press. Stillwater, MN, USA.
Karanth, K.U. and Nichols, J.D. 1998. Estimating tiger densities in India from camera trap
data using photographic captures and recaptures. Ecology 79:2852-2862.
Karanth, K.U. and Nichols, J.D. 2000. Ecological status and conservation of tigers in India.
Final Technical Report to the US Fish and Wildlife Service (Division of International
 Monitoring Tigers and Prey: Conservation Needs and Managerial Constraints
Conservation), Washington, DC, and Wildlife Conservation Society, New York. Centre
for Wildlife Studies, Bangalore, India.
Karanth, K.U., Nichols, J.D., Seidensticker, J., Dinerstein E., Smith, J.L.D., McDougal C.,
Johnsingh, A.J.T., Chundawat, R.S. and Thapar, V. Science deficiency in conservation
practice: The monitoring of tiger populations in India. (In prep.).
Karanth, K.U. and Sunquist, M.E. 1992. Population structure, density and biomass of large
herbivores in the tropical forests of Nagarahole, India. Journal of Tropical Ecology
8:21-35.
Lancia, R.A., Nichols, J.D. and Pollock, K.N. 1994. Estimation of number of animals in
wildlife populations. Pages 215-253 in Research and management techniques for
wildlife and habitats (Ed: T. Bookhout ). The Wildlife Society, Bethesda, MD,
USA.
Nichols, J.D. 1992. Capture-Recapture models: Using marked animals to study population
dynamics. Bioscience 42:94-102.
Rabinowitz, A. 1997. Wildlife field research and conservation training manual. Wildlife
Conservation Society, New York, NY, USA.
Seber, G.A.F. 1982. The estimation of animal abundance and related parameters. Second
edition. MacMillan, New York, NY, USA.
Thompson, S.K. 1992. Sampling. Wiley, New York, NY, USA.
Thompson, W.L., White, G.C. and Gowan, C. 1998. Monitoring vertebrate populations.
Academic Press, New York, USA.
Williams, B.K., Nichols, J.D. and Conroy, M.J. 2002. Analysis and management of animal
populations. Academic Press, San Diego, CA, USA.
8 9
CHAPTER 2
ECOLOGY OF THE TIGER: IMPLICATIONS FOR
POPULATION MONITORING
K. Ullas Karanth1 and Raghunandan S. Chundawat2
Wildlife Conservation Society (International Programs), Bronx, New York, NY-10460-1099, USA.
1
Email: <ukaranth@wcs.org>
Wildlife Institute of India, P.B. 18, Chandrabani, Dehra Dun, Uttaranchal-248 001, India.
2
Email: <pnptiger@mantraonline.com>
INTRODUCTION
In the previous chapter, we saw how logistical and social factors, such as availability of
technical skills, adequate manpower, and material resources must be considered carefully
before establishing monitoring programs for tigers or their prey. However, apart from such
factors, the ecology of the tiger also significantly influences the monitoring program. Therefore,
monitoring of tigers and their prey must also consider how ecological factors affect population
assessment methods. In this chapter we provide a quick overview of the findings from scientific
research on tigers, focusing on aspects of tiger biology that have a bearing on population
assessment and monitoring.
Many naturalists and hunters have recorded anecdotal accounts of tigers in tropical Asia
during the past two centuries (Karanth 2001). For example, during the 20th century, important
accounts of tiger behavior were provided by Brander (1923), Corbett (1944), Singh (1984)
and Thapar (1989) in tropical Asia. Although qualitative in nature, these accounts provided
us valuable insights.
Scientific studies of tigers within the framework of quantitative wildlife biology were pioneered
by George Schaller (Schaller 1967). Further scientific advances were made in the 1973-1985
period through the radio telemetry studies at Chitwan, Nepal, under the Smithsonian Tiger
Ecology Project (Seidensticker 1976; McDougal 1977; Sunquist 1981; Smith et al. 1987;
Smith 1993). In the1990’s long-term ecological studies of tigers were begun by Ullas Karanth
and his collaborators (Karanth and Sunquist 1992, 1995, 2000; Karanth and Nichols 1998,
2000; Karanth and Stith 1999; Karanth et al. 1999) and Raghunandan Chundawat and
colleagues (Chundawat et al. 1999) in India, and by Dale Miquelle and partners (Miquelle
et al. 1999) in Russia. These long-term studies have employed radio telemetry, camera trapping,
diet analyses and prey density estimation. They provide a rigorous basis for examining the
issues of monitoring tigers and their prey (Karanth 2001).
 Ecology of the Tiger: Implications for Population Monitoring Ecology of the Tiger: Implications for Population Monitoring

The post-1980 period has witnessed a remarkable blooming of population assessment

methodologies in the wildlife sciences (Burnham et al. 1980; Seber 1982; Nichols 1992;
Thompson 1992; Buckland et al. 1993, 2001; Lancia et al. 1994; Thompson et al. 1998;
Williams et al. 2002). This growth was nurtured by an increased involvement of mathematicians
and biostatisticians in the field of population ecology. The phenomenal growth of the
information technology industry has also enabled biologists to overcome earlier computational
limitations. Software are now readily available that use complex algorithms and iterative
procedures needed to implement biologically realistic population estimation protocols (Nichols
1992).

A major goal of this manual is to integrate these modern methods of population assessment

Figure 2.1 - Tiger social organization pattern at the landscape level.

and monitoring with our current knowledge of tiger ecology. Therefore, in this chapter, we
summarize recent findings on tiger biology, and describe how these impinge on population
monitoring issues.

TIGER SOCIAL ORGANIZATION AND LAND TENURE SYSTEM

A population of wild tigers includes individuals belonging to both sexes and to different age-
classes. Here, following Karanth and Stith (1999), we have classified tigers into four age
classes (demographic stages) as: Cubs (< 12 month old tigers), juveniles (12-24 month old
pre-dispersal tigers), transient ‘floaters’ (>24 month old, dispersing tigers that do not breed
or hold stable home ranges) and resident breeders (tigers that maintain stable ranges and
reproduce).

Tiger society revolves around adult breeding females. These tigresses maintain relatively
fixed home ranges within which they raise cubs. Normally, a tigress acquires her own home
range and starts breeding at 3-4 years of age. Thereafter, she has tenure of about 5-7 years,
before she loses her range to a more vigorous competitor. The degree of overlap between
neighboring female ranges (or absence of such overlap) appears to vary among areas depending
on prey density and other ecological factors (Sunquist 1981; Smith et al. 1987; Miquelle
et al. 1999; Karanth and Sunquist 2000).

Adult male tigers have larger ranges that overlap ranges of several breeding females,
the average being about three (Figure 2.1). The extent of overlap between male ranges appears
to vary, depending on factors that are as yet unclear. The tenures of breeding males appear to
be significantly shorter (2-4 years) than the 6-10 year tenures of females (Sunquist 1981;
Smith 1993).

10 11
 Ecology of the Tiger: Implications for Population Monitoring
A tigress having favorable conditions within her range produces a litter of 3-4 cubs (following
a gestation of about 105 days), once every 2-3 years. She raises the cubs on her own. At
about 18-24 months age, the juveniles are evicted by their mother from her range and become
transients. Therefore, in addition to resident breeders and cubs, a tiger population may also
have pre-dispersal juveniles and dispersing transients (floaters) (Smith 1993; Kenny et al.
1995; Karanth and Stith 1999).
During their wanderings, the transient floaters move over dozens of kilometers and range
across several breeder territories, constantly looking for vacancies to settle down. The presence
of floaters appears to be tolerated to some extent by their mothers, and by the specific individual
male tigers that sired them. When a breeder becomes old, weak or injured, its tenure is ended
by a more vigorous rival (either a neighbor or a transient) who takes over its range. A small
segment of the transient population may also comprise of evicted breeders, although such
animals may be short-lived.
Tigers seek or avoid social contact, depending on the context, using a complex communication
system based on scent marking, visual marking and vocalizations (Smith et al. 1989). Such
tiger signs include: Scent sprays from anal glands, scent deposits from digital glands, tracks
and scrape marks, scat deposits, claw marks on trees and vocalizations of several kinds.
Using these mechanisms tigers are able to maintain social contact even at the relatively low
densities at which they live (Karanth and Nichols 1998, 2000). However, as far as monitoring
by humans is concerned, tigers are extremely secretive animals: Except for a few individuals
habituated to tourists, throughout their range they are difficult animals to find and observe.
ECOLOGICAL DETERMINANTS OF TIGER DENSITY
Over most of their range, tigers coexist with other predatory carnivores such as leopards,
dholes or wolves. The relative numbers (densities) of different predator species within such
guilds appear to be primarily determined by the relative abundance of different size classes of
prey species in the assemblage (Karanth and Sunquist 1995; Karanth and Nichols 1998).
Furthermore, densities of tigers appear to be primarily a function of prey densities (Schaller
1967; Sunquist 1981; Seidensticker and McDougal 1993; Karanth 1995; Karanth and Sunquist
1995; Karanth and Nichols 1998; Chundawat et al. 1999; Sunquist et al. 1999).
It is clear from basic principles of animal energetics (Eisenberg 1980) that tiger’s principal
prey must be large (>20 kg). For example, just as lions cannot live off the high biomass of
termites in African grasslands, tigers cannot attain high densities in the absence of enough
ungulate prey, even if smaller prey are abundant (Schaller 1967; Sunquist 1981; Karanth and
12
Ecology of the Tiger: Implications for Population Monitoring
Sunquist 1995; Sunquist et al. 1999). Recent studies that estimated tiger and prey abundance
using rigorous methods (Karanth and Nichols 1998, 2000) clearly show a strong positive
relationship between prey abundance and tiger densities (Figure 2.2).
Densities of principal prey species influence tiger densities in several ways. As prey densities
decline, breeding female ranges become larger, dramatically reducing the number of such
females that an area can support. For instance, the size of female home ranges in productive
South Asian forests and grassland is 10-20 km2, whereas in the Russian Far East it is as large
as 200-400 km2 (Sunquist 1981; Miquelle et al. 1999; Karanth and Sunquist 2000). High
prey densities appear to permit the area to support higher number of transient animals also
(Karanth and Nichols 1998, 2000). Because cub and juvenile survival rates are higher when
prey availability is higher, the numbers of tigers in these two demographic stages are also
higher. While other habitat-related or managerial factors also influence tiger density at a
given site, prey abundance appears to be the primary ecological determinant in most places.
Because tiger densities are strongly correlated with prey densities, regardless of the survey
method employed, our encounter rates with tigers or their sign (detections of tigers or their
signs, photo trap success rates, etc.) will also be strongly influenced by prey densities.
Therefore, in areas with high prey densities, because of potentially larger sample sizes, more
reliable sign-based indices of relative abundances of tigers or prey can be derived.
SURVIVAL, MORTALITY AND POPULATION DYNAMICS
Tigers in all demographic stages die from a variety of causes. Like other cats, tigers too
exhibit infanticidal behavior: A ‘new’ male tries to kill the cubs of the former resident breeder
whom he replaces. Adult males appear to tolerate only their own offspring during occasional
encounters. Cubs may also die due to neglect or inexperience on the part of their mother.
Other factors that cause heavy cub mortalities are: Starvation, floods, forest fires, other
predator species, and, human persecution - either directly, or indirectly through the killing of
the mother.
Juvenile tigers (1-2 years age) die from starvation, hunting injuries and intra-specific
aggression, although their mortality rates are lower than those of cubs. During dispersal,
tigers move back and forth through larger ecological units, traveling across hundred kilometers
or more, crisscrossing the boundaries of breeder territories, nature reserves and even states
or countries (Smith 1993). During the dispersal phase, tigers suffer high mortality rates
from starvation, intra-specific aggression and human persecution as they range across
vast landscapes.
13
 Ecology of the Tiger: Implications for Population Monitoring
Figure 2.2 - A tiger ‘crops’ 50 animals every year from a base population of 500 prey animals.
14
Ecology of the Tiger: Implications for Population Monitoring
The approximate age-specific annual survival rates for tigers in productive habitats (Kenny et al.
1995; Karanth and Stith 1999; K.U. Karanth and J.D. Nichols unpublished data) are: Cubs=60%;
Juveniles=90%; Female Transients=70%; Male Transients=65%; Breeding Females=90%;
Breeding Males=80%. These rates are much lower (mortality rates higher) in sub-optimal habitats.
It should be noted that such high natural mortality rates are not necessarily a cause for alarm;
they are naturally complemented by the tiger’s high reproductive potential and recruitment
rates.
Demographic simulations (Kenny et al. 1995; Karanth and Stith 1999) suggest that cubs less
than a year old may comprise roughly 25% of a healthy tiger population. The average
reproductive potential of female breeders in prey-rich tiger habitats may exceed one cub per
female per year. In other words, with no constraints on prey availability and habitat, a tigress
replaces herself several times during her own lifetime, producing a large ‘surplus’ of tigers
that must disperse out of the area. Consequently, the high mortality rates do not depress tiger
population densities, as long as the reproductive rates are also naturally high.
Because of these high mortality and reproduction rates, tiger populations experience relatively
high turnover of individuals. In addition, factors such as transient behavior, dispersal
movements and immigration also add to the turnover rate. As a result of this high turnover in
every wild tiger population, a large proportion of animals present each year comprise of
‘new’ individuals.
EFFECT OF ENVIRONMENTAL FACTORS ON MONITORING
Tigers occur in a variety of environments: Temperate boreal and coniferous forests in Russia;
subtropical forests in China, Nepal, Bhutan, and India; mangrove forests in India and
Bangladesh; alluvial grasslands in India and Nepal; and tropical wet-evergreen, moist-
deciduous and dry-deciduous forests all over tropical Asia (Wikramanayake et al. 1999).
The environmental variables associated with the different habitat types will also strongly
influence encounter rates of survey teams with tiger signs.
In the Russian Far East and adjoining areas in China, fresh snowfall provides an ideal substrate
for retaining tiger tracks during winter. Similarly, the perennially moist substrate in the
mangrove forests of Sundarbans of India and Bangladesh also permit retention of track prints.
In contrast, almost everywhere else, tiger tracks are readily seen only on trails, riverbeds or
roads where the substrate is suitable due to natural factors or vehicular traffic. Because tigers
prefer natural trails and man-made roads as travel routes, their tracks and other signs are
more likely to be encountered on them.
15
 Ecology of the Tiger: Implications for Population Monitoring
However, over most of tiger range - particularly in forests with hard soil or leaf-litter covered
substrates - it is very difficult to find and record tiger signs such as tracks, scats and scrapes.
Although tiger scats can be detected on substrates where tracks are not visible, these may
decompose rapidly due to weather or insect activity. Light rains create conditions conducive
to finding tiger signs during surveys, but heavy rainfall will wash the signs away.
TIGER ECOLOGY: CHALLENGES TO MONITORING
Most aspects of tiger ecology pose serious challenges to those who try to monitor tiger
populations. Tigers are usually shy and avoid human contact. Their secretive nature, preference
for dense cover and largely nocturnal ranging patterns make them difficult to observe during
surveys. Even tiger signs such as tracks and scats are dispersed widely and difficult to detect
over larger regions, although they may be quite abundant at specific sites within major protected
areas. Tiger tracks show up readily where the substrate is snow, mud or fine alluvial soil,
particularly on roads and riverbeds. However, over large parts of tiger range in tropical Asia,
substrate conditions are unsuitable and road densities low, making sign surveys difficult.
Another major problem is posed by the normal ranging behavior of tigers. Even resident
breeders can move over 10-15 km per day, leaving their signs at widely separated localities
even within a short span of a week or less. The problem is even worse with transient tigers
that move over tens or even hundreds of kilometers. They leave signs while moving back and
forth across human constructs such as boundaries of census blocks, parks, regions or even
States. In addition, despite their precarious status, tigers are distributed over wide areas in
tropical Asia, with the potential tiger habitat estimated to be around 1,200,000 km 2
(Wikramanayake et al. 1999).
However, the most difficult challenge to monitoring is the basic fact that, being large-bodied
carnivores, tigers live at extremely low densities. Even in the best sites, densities are as low as
10-20 tigers/100 km2. In areas with low prey densities, a situation that prevails over 90%
of tiger range, densities can be as low as 0.5-1.0 tigers/100 km2 (Karanth and Nichols
1998, 2000).
The combination of all the above factors - low densities, secretiveness, wide-ranging behavior,
extensive distribution and the low probabilities of detecting tiger sign – poses serious
methodological problems. Collection of quantitative data becomes problematic because
of small sample sizes and logistical or practical constraints faced by field survey teams.
Therefore, typically, even results from simple distribution surveys of tiger sign tend to lack
robustness.
16
Ecology of the Tiger: Implications for Population Monitoring
HOW TO MONITOR TIGERS
Tiger conservation and management can be viewed as occurring at three different scales or
levels of resolution. Monitoring needs and objectives differ among these three levels and lead
to different kinds of monitoring programs with differing methods.
Tigers and their principal prey are landscape animals. As habitats get fragmented and come
under pressure, local populations may be extirpated. The distributional range of tigers may
contract and shrink. On the other hand, if conservation measures are successful, habitat
fragments may be reconnected, creating larger populations. Dispersal into newly improved
habitats can establish new populations. Therefore, at the national or regional level it is critical
for managers, biologists and conservationists to accurately document, on a continuous basis,
the presence and absence of tigers and principal prey species.
At the level of individual reserves, managers, researchers or conservationists may have to
keep track of population trends of tigers and prey, to evaluate the utility of various conservation
interventions. Although desirable, it may not always be possible to obtain rigorous estimates
of numbers or densities of tigers. However, many management needs can be met by reliable
indices of population trends. Such indices of relative abundance may allow comparison of
population densities between two sites, or over time at the same site.
At specific study sites, where the primary goal is to understand tiger ecology at a fine-grained
level, parameters such as tiger population size, absolute density, and rates of survival, mortality,
immigration and dispersal may have to be estimated. In this case, there is no escape from the
need for employing the most advanced equipment, techniques and skilled personnel. Such
fine-grained data cannot be obtained for tigers employing only simple techniques and basic
equipment. Obtaining such fine-grained data is primarily the goal of wildlife biologists.
THE NEED FOR A UNIFIED FRAMEWORK FOR MONITORING
The three classes of goals listed above are distinct and are often pursued by different agencies
or individuals. However, they are mutually contributory. It is unwise for the practitioners of
each to go their separate ways. The underpinning of all tiger population monitoring must be
provided by sound science. While we agree that the methods applied must match the resources
and skills available, lack of resources or training should not be used as an excuse to practice
substandard science. In other words, having worthless data is worse than having no data at all
(Karanth 1999).
17
 Ecology of the Tiger: Implications for Population Monitoring
It is clear that the methods employed to meet data collection needs at different scales and
levels of resolution are likely to be quite different in terms of the sampling design, effort,
equipment used and the kinds of personnel skills required. However, regardless of the variables
involved in a particular situation, the monitoring methods must be based on good science.
This means they should meet general criteria such as a sound theoretical basis, a history of
empirical validation of assumptions and results, and a track record of being refined through
the universally accepted process of scientific peer review.
Considering the radically different needs of monitoring, the variety of field situations and the
differences in available manpower, equipment and resources, it is clear that an array of different
methods – rather than any single approach – is required. While these tiger monitoring methods
must work in different ecological, social and technological contexts, they should also be
based on a sound scientific framework. A unified, rigorous conceptual framework that
accommodates an array of valid methods for monitoring tigers and their prey is provided in
Chapter 3.
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Buckland, S.T., Anderson, D.R., Burnham, K.P. and Laake, J.L. 1993. Distance sampling:
Estimating abundance of biological populations. Chapman and Hall, London, UK.
Buckland, S.T., Anderson, D.R., Burnham, K.P., Laake, J.L., Borchers, D.L. and Thomas,
L. 2001. Introduction to distance sampling. Oxford University Press, Oxford, UK.
Burnham, K.P., Anderson, D.R. and Laake, J.L. 1980. Estimation of density from line transect
sampling of biological populations. Wildlife Monographs 72:1-202.
Chundawat, R.S., Gogate, N. and Johnsingh, A.J.T. 1999. Tigers in Panna: Preliminary
results from an Indian tropical dry forest. Pages 123-129 in Riding the tiger: Tiger
conservation in human-dominated landscapes (Eds: J. Seidensticker, S. Christie
and P. Jackson). Cambridge University Press, Cambridge, UK.
Corbett, J. 1944. Man-eaters of Kumaon. Oxford University Press, London, UK.
Eisenberg, J.F. 1980. The density and biomass of tropical mammals. Pages 34-55 in
Conservation Biology: An evolutionary-ecological perspective (Eds: M.E. Soulé
18
Ecology of the Tiger: Implications for Population Monitoring
and B.A.Wilcox). Sinauer Associates, Sunderland, MA, USA.
Karanth, K.U. 1995. Estimating tiger populations from camera-trap data using capture-
recapture models. Biological Conservation 71:333-338.
Karanth, K.U. 1999. Counting tigers, with confidence. Pages 350-353 in Riding the tiger:
Tiger conservation in human-dominated landscapes (Eds: J. Seidensticker, S. Christie
and P. Jackson). Cambridge University Press, Cambridge, UK.
Karanth, K.U. 2001. The way of the tiger: Natural history and conservation of the endangered
big cat. Voyageur Press. Stillwater, MN, USA.
Karanth, K.U. and Nichols, J.D. 1998. Estimating tiger densities in India from camera trap
data using photographic captures and recaptures. Ecology 79:2852-2862.
Karanth, K.U. and Nichols, J.D. 2000. Ecological status and conservation of tigers in India.
Final Technical Report to the US Fish and Wildlife Service (Division of International
Conservation), Washington, DC, and Wildlife Conservation Society, New York. Centre
for Wildlife Studies, Bangalore, India.
Karanth, K.U. and Stith, M.B. 1999. Prey depletion as a critical determinant of tiger population
viability. Pages 100-113 in Riding the tiger: Tiger conservation in human-dominated
landscapes (Eds: J. Seidensticker, S. Christie and P. Jackson). Cambridge University
Press, Cambridge, UK.
Karanth, K.U. and Sunquist, M.E. 1992. Population Structure, density and biomass of large
herbivores in the tropical forests of Nagarahole, India. Journal of Tropical Ecology
8:21-35.
Karanth, K.U. and Sunquist M. E. 1995. Prey selection by tiger, leopard and dhole in tropical
forests. Journal of Animal Ecology 64:439-450.
Karanth, K.U. and Sunquist, M.E. 2000. Behavioral correlates of predation by tiger, leopard
and dhole in Nagarahole, India. Journal of Zoology (London). 250:255-265.
Karanth, K.U., Sunquist, M.E. and Chinnappa, K.M. 1999. Long term monitoring of tigers:
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19
 Ecology of the Tiger: Implications for Population Monitoring Ecology of the Tiger: Implications for Population Monitoring

Kenny, J.S., Smith, J.L.D., Starfield, A.M. and McDougal, C. 1995. The long-term effects Smith J.L.D., McDougal C. and Miquelle, D. 1989. Scent marking in free ranging tigers,
of tiger poaching on population viability. Conservation Biology 9:1127-1113. Panthera tigris. Animal Behavior 37:1-10.

Lancia, R.A., Nichols, J.D. and Pollock, K.N. 1994. Estimation of number of animals in
wildlife populations. Pages 215-253 in Research and management techniques for
wildlife and habitats. (Ed: T. Bookhout ). The Wildlife Society, Bethesda, MD,
USA.
McDougal, C. 1977. The face of the tiger. Rivington Books, London, UK.
Miquelle, D.G., Smirnov, E.N., Merrill, T.W., Myslenkov, A.E., Quigley, H.B., Hornocker,
M.G. and Schleyer B. 1999. Hierarchical spatial analysis of Amur tiger relationships
to habitat and prey. Pages 71-99 in Riding the Tiger: Tiger conservation in human
dominated landscapes (Eds: J. Seidensticker, S. Christie and P. Jackson). Cambridge
University Press, Cambridge, UK.
Nichols, J.D. 1992. Capture-Recapture models: Using marked animals to study population
dynamics. Bioscience 42:94-102.
Schaller, G.B. 1967. The deer and the tiger. University of Chicago Press, Chicago, USA.
Seber, G.A.F. 1982. The estimation of animal abundance and related parameters. Second
edition. MacMillan, New York, NY, USA.
Seidensticker, J. 1976. On the ecological separation between tigers and leopards. Biotropica
8:225-234.
Seidensticker, J. and McDougal C. 1993. Tiger predatory behavior, ecology and conservation.
Symposium of the Zoological Society of London 65:105-125.
Sunquist M.E. 1981. Social organization of tigers (Panthera tigris) in Royal Chitawan National
Park, Nepal. Smithsonian Contributions to Zoology 336:1-98.
Sunquist M.E., Karanth K.U. and Sunquist F. 1999. Ecology, behavior and resilience of the
tiger and its conservation needs. Pages 5-18 in Riding the tiger: Tiger conservation
in human-dominated landscapes (Eds: J. Seidensticker, S. Christie and P. Jackson).
Cambridge University Press, Cambridge, UK.
Thapar, V. 1989. Tigers: The secret life. Elm Tree books, London, UK.
Thompson, S.K. 1992. Sampling. Wiley, New York, NY, USA.
Thompson, W.L., White, G.C. and Gowan, C. 1998. Monitoring vertebrate populations.
Academic Press, New York, NY, USA.
Wikramanayake E.D., Dinerstein E., Robinson J.G., Karanth K.U., Rabinowitz A., Olson
D., Matthew T., Hedao P., Conner M., Hemley G. and Bolze D. 1999. Where can
tigers live in the future? A framework for identifying high priority areas for the
conservation of tigers in the wild. Pages 255-272 in Riding the tiger: Tiger
conservation in human-dominated landscapes (Eds: J. Seidensticker, S. Christie
and P. Jackson). Cambridge University Press, Cambridge, UK.
Williams, B.K., Nichols, J.D. and Conroy, M.J. 2002. Analysis and management of animal
populations. Academic Press, San Diego, CA, USA.

Singh, A. 1984. Tiger! Tiger! Jonathan Cape. London, UK.

Smith J.L.D. 1993. The role of dispersal in structuring the Chitwan tiger population. Behavior
124:165-195.

Smith J.L.D., McDougal C., and Sunquist M.E. 1987. Female land tenure system in tigers.
Pages 97-109 in Tigers of the World: The biology, biopolitics, management and
conservation of an endangered species (Eds: R.L. Tilson and U.S. Seal). Noyes
Publications, Park Ridge, NJ, USA.

20 21
 CHAPTER 3

POPULATION MONITORING: A CONCEPTUAL FRAMEWORK

James D. Nichols1 and K. Ullas Karanth2

United States Geological Survey, Patuxent Wildlife Research Center, Laurel, MD-20708-4017, USA.
1

Email: <jim_nichols@usgs.gov>

Wildlife Conservation Society (International Programs), Bronx, New York, NY-10460-1099, USA.
2

Email: <ukaranth@wcs.org>

INTRODUCTION

“Monitoring” of animal populations can be defined as the estimation of absolute or relative

abundance for the purpose of drawing inferences about variation in abundance of animals
BLANK over space and/or time. For example, we might focus on a single time period and ask whether
animal abundance varies among different locations at that time. Such inferences are useful
for assessing the distribution patterns of animals as well as for addressing questions about
the relationship between animal abundance and such factors as habitat or management actions.
We might also focus on a single location and ask whether animal abundance varies over time.
Rates of temporal change are sometimes referred to as “trends”, and some workers view
monitoring as restricted to such assessments of temporal change. In this manual, we discuss
different approaches for use in abundance estimation and monitoring of populations of tigers
and their prey in the forests of tropical Asia.

Although we will present different monitoring methods and approaches, we believe that it is
useful to emphasize the common conceptual basis that underlies all such methods. This
conceptual framework clarifies the relationship among these seemingly disparate methods
and provides a basis for considering and developing modifications and new methods. In
particular, this framework establishes the relationship between indices and abundance estimates
and should permit informed decisions about which of these approaches to follow in different
situations.

THE STATISTICAL FRAMEWORK

Basic Problems in Counting Animals

Virtually all inferences about animal populations are based on count statistics. In many

23
 Population Monitoring: A Conceptual Framework
cases, count statistics are provided by direct counts of animals themselves. For example, we
might count the number of chital deer observed while walking on a line transect or the number
of tigers identified (caught) by camera trapping. In other situations, count statistics are based
on animal sign such as ungulate tracks, tiger pugmarks, tiger scats, and dung of prey species.
Two basic problems confront biologists and managers who would like to use such count
statistics to estimate and draw inferences about animal population size: Observability and
spatial sampling (see Nichols 1992; Thompson 1992; Lancia et al. 1994; Skalski 1994;
Nichols and Conroy 1996; Williams et al. 2002). With respect to direct counts of animals,
observability refers to the usual inability to detect and enumerate all animals, regardless of
the sampling or survey method being used. With respect to indirect evidence of animal presence
such as tracks, pugmarks, scat and dung, observability refers more generally to an inequality
between the count statistic and the true number of animals. Spatial sampling, on the other
hand, refers to the fact that we are frequently interested in areas so large that we are unable to
obtain our count statistics over the entire area. Instead, we must select smaller areas thought
to be representative of the entire area, with the idea that we will try to use counts on these
sampled areas to draw inferences about the number of animals in the entire area.
Observability
We consider observability by first defining the following quantities obtained on a sample area
(or on the entire area, if we assume there is no need to sub-sample areas):
C = count statistic, or number of animals (or indirect sign) counted;
N = abundance, or true number of animals;
p = proportionality constant relating the count statistic and abundance.
In the case of direct counts of animals, p reflects the probability that an animal in the sampled
area is counted (the probability that a member of N appears in the count statistic; p can also
be viewed as the expected proportion of animals appearing in the count statistic). The following
expression shows the relationship between the count statistic and abundance:
E(C) = Np (3.1)
where E(C) denotes the expected value (or expectation) of C.
C is a random variable and can assume different values each time a count is made. E(C) can
be viewed as the average value of C that would be obtained if the count statistic could be
collected a large number of times in the same exact sampling situation with the same N. In
24
Population Monitoring: A Conceptual Framework
cases where C represents some count of animal sign (or of something other than the animals
themselves), equation 3.1 still provides a reasonable model. However, in the case of animal
sign, p should not be thought of as a detection probability but simply as a coefficient relating
N and C.
Estimation of abundance is based on the estimation of p in equation 3.1. If we are able to
estimate the p associated with a particular count statistic (denote this estimate as pˆ where
the “hat” denotes an estimate), the abundance can be estimated as:
C
Nˆ = (3.2)
pˆ
The estimator in equation (3.2) is very general, as virtually all population estimation methods
(Seber 1982) for a single location can be written in this general form. For example, the count
statistic under distance sampling (Buckland et al. 1993, 2001) is the number of animals
observed and counted (e.g., along a line transect), and the perpendicular distances of these
observations to the transect line are used to estimate the detectability function and, hence, p.
The count statistic under capture-recapture sampling (Otis et al. 1978; Seber 1982; White
et al. 1982; Pollock et al. 1990) is the number of different animals caught, and the patterns of
capture and recapture for individual animals are modeled in order to estimate p.
As a numerical example of the rationale underlying equation 3.2, assume that we count 20
deer in an area and estimate a corresponding detection probability of pˆ = 0.25 ; that is we
estimate that we detected about 25% of the animals when we conducted our counts. Our
abundance estimate is then obtained as Nˆ =20/0.25 = 80. This estimate is intuitively reasonable
in that we estimate that we detect approximately one of every four deer, so our estimated
abundance is four times the number of animals counted. Perhaps the most important
consideration resulting from equations 3.1 and 3.2 is that the count statistic itself does not
permit unambiguous inference about abundance. Instead, such inference requires information
about the detection probability associated with the count statistic.
Spatial Sampling
Typically, we cannot survey an entire area of interest, so we must select sample areas thought
to be representative of the entire area. These sample areas will represent some fraction, a, of
the total area. Unlike the situation where the fraction of animals present in a sampled area
that is counted (p) must be estimated, the spatial sampling fraction (a ) is typically known
and requires no estimation. Define Nˆ ′ to be the estimated abundance of animals in sampled
areas representing fraction a of the total area of interest (one means of achieving
representativeness is through simple random sampling). Then abundance for the
25
 Population Monitoring: A Conceptual Framework Population Monitoring: A Conceptual Framework

entire area of interest can be estimated as: is estimated using replication and will depend on the spatial distribution of animals over the
study area and the selected sampling design. This component is small when animals are
Nˆ ′ evenly distributed over space and large when animals are clumped. There is also a variance
Nˆ = (3.3)
α component associated with the estimation of p, var( pˆ ). In general, smaller variances of
abundance estimates, var( Nˆ ), result from smaller variance components, var(C) and var( pˆ ),
i.e., we simply divide the estimated abundance for the sampled locations by the fraction of the and from larger values of detection probability, p, and proportion of the area that is
entire area represented by those locations. sampled, a.

As a numerical example, assume that we have randomly, or at least representatively, sampled Discussion
several locations representing 10% of the entire area of interest (a = 0.10), and that we have
obtained an estimate of 80 deer on these locations. Then the population estimate for the entire Finally, we emphasize that equations 3.1 – 3.4 do not represent a specific estimation method
area is computed as Nˆ = 80 / 0.10 = 800 deer. for animal abundance. Instead, they provide a conceptual framework for thinking about
abundance estimation problems. Indeed, all of the specific abundance methods presented in
Canonical Estimator the reviews by Seber (1982), Lancia et al. (1994) and Williams et al. (2002) can be viewed as
special cases that fall within this general framework. In addition to providing a unifying
We then combine the above solutions to the problems of observability and spatial sampling conceptual approach for viewing these established methods, we believe that this framework
(equations 3.2 and 3.3) into a single, general estimator: provides a useful way to think about abundance estimation that can lead to new methods
tailored to the logistical and ecological specifics of new estimation problems.
C′
Nˆ = (3.4)
pˆ α
REFERENCES
where C ′ is the count statistic on sampled areas and pˆ is the estimated detection probability,
assumed equal for all samples in this expression (this need not be assumed in general; see
Buckland, S.T., Anderson, D.R., Burnham, K.P. and Laake, J.L. 1993. Distance sampling:
Thompson 1992; Skalski 1994). We thus estimate population size by dividing the count
Estimating abundance of biological populations. Chapman and Hall, London, UK.
statistic by both the estimated fraction of the animals on the sampled area(s) that were detected
( pˆ ), and the proportion of the total area from which the count statistic was taken (a). We
Buckland, S.T., Anderson, D.R., Burnham, K.P., Laake, J.L. Borchers, D.L. and Thomas, L.
refer to expression 3.4 as the canonical estimator for abundance.
2001. Introduction to distance sampling. Oxford University Press, Oxford, UK.
For example, assume that we count 20 deer (C ′ = 20 ) on sample areas representing 10% of
Lancia, R.A., Nichols, J.D. and Pollock, K.N. 1994. Estimation of number of animals in
an area of interest, so a = 0.10. Further assume that we are able to estimate a detection
wildlife populations. Pages 215-253 in Research and management techniques for
probability of 0.25 ( pˆ = 0.25). Then we estimate abundance as:
wildlife and habitats. (Ed: T. Bookhout ). The Wildlife Society, Bethesda, MD, USA.
20
Nˆ = = 800 deer.
(0.10) (0.25) Nichols, J.D. 1992. Capture-recapture models: Using marked animals to study population
dynamics. BioScience 42:94-102.
The exact form of the variance of the canonical estimator depends on the spatial sampling
design and on variation in detection probability over the different sampled locations (e.g., see Nichols, J.D. and Conroy, M.J. 1996. Estimation of mammal abundance: Introduction.
Thompson 1992; Skalski 1994). However, it is useful to consider the general components of Pages 177-179 in Measuring and monitoring biological diversity: Standard methods
the variance estimator and their general effects on the magnitude of the variance. One component for mammals (Eds: D.E. Wilson, F.R. Cole, J.D. Nichols, R. Rudran and M.S. Foster).
is the variance of the count statistic among the different sampled areas, var(C). This component Smithsonian Institution Press, Washington, DC, USA.

26 27
 Population Monitoring: A Conceptual Framework CHAPTER 4

Otis, D.L., Burnham, K.P., White, G.C. and Anderson. D.R. 1978. Statistical inference from STATISTICAL CONCEPTS: ASSESSING SPATIAL DISTRIBUTIONS
capture data on closed animal populations. Wildlife Monographs 62:1-135.

Pollock, K.H., Nichols, J.D., Brownie, C. and Hines, J.E. 1990. Statistical inference for James D. Nichols1 and K. Ullas Karanth2
capture-recapture experiments. Wildlife Monographs 107:1-97.
United States Geological Survey, Patuxent Wildlife Research Center, Laurel, MD-20708-4017, USA.
1

Email: <jim_nichols@usgs.gov>
Seber, G.A.F. 1982. The estimation of animal abundance and related parameters. MacMillan,
New York, NY, USA. Wildlife Conservation Society (International Programs), Bronx, New York, NY-10460-1099, USA.
2

Email: <ukaranth@wcs.org>

Skalski, J.R. 1994. Estimating wildlife populations based on incomplete area surveys. Wildlife
Society Bulletin 22:192-203.
PRESENCE-ABSENCE SURVEYS
Thompson, S.K. 1992. Sampling. John Wiley and Sons, New York, NY, USA.
It is frequently easier to survey an area to determine whether or not tigers or prey animals are
present, than to collect the sorts of statistics needed to estimate absolute abundance or relative
White, G.C., Anderson, D.R., Burnham, K.P. and Otis, D.L. 1982. Capture-recapture and
abundance. For certain large-scale surveys directed at geographic distribution of tigers or
removal methods for sampling closed populations. Los Alamos National Laboratory
prey species, it is reasonable to consider use of presence-absence surveys. We focus this
Publication LA-8787-NERP. Los Alamos, NM, USA.
discussion on tiger populations, but note that it is equally reasonable to apply these methods
to the tiger’s prey species. The design of a presence-absence survey depends on the study
Williams, B.K., Nichols, J.D. and Conroy, M.J. 2002. Analysis and management of animal
objectives. The objective for such surveys is typically one of the following:
populations. Academic Press, San Diego, CA, USA.
1. Estimation of proportion of area occupied by tigers.
2. Mapping of tiger distribution.

Estimating the Proportion of Area Occupied

Design of a survey to estimate the proportion of area occupied first requires division of the
area of interest into grid cells (e.g., 10 km2), or some other form of spatial sampling unit.
Define the quantity of interest as:

R
ψ =
S
where ψ = proportion of area occupied, R = number of grid cells or sample plots containing
tiger sign, and S = number of grid cells in the entire area. The proportion area occupied can
then be estimated as:

Rˆ
ψˆ = (4.1)
S
where S is known, but R (and thus ψ ) is not known and must be estimated.

28 29
 Statistical Concepts: Assessing Spatial Distributions Statistical Concepts: Assessing Spatial Distributions

As in most animal sampling situations (see Chapter 3), there are two sources of variation to
consider in the estimation of R and ψ : Spatial sampling and observability. Spatial sampling
is not an issue if all S cells are surveyed. Possible approaches to spatial sampling when all
cells are not surveyed include simple random sampling, stratified random sampling, and adaptive
cluster sampling. Simple random sampling requires random selection of s cells for survey
from the total number of S potential cells. Stratified random sampling first involves division
of the entire area of interest into strata based on a factor such as habitat or expected tiger
density. Then, simple random sampling of cells is used within each stratum. Allocation of
samples to the different strata may be proportional to stratum size, proportional to expected
tiger density, or according to some other scheme (Thompson 1992). Adaptive cluster sampling
begins with simple random sampling. The second step involves sampling of grid cells that
border those cells in the initial sample that contained sign. The final step involves sampling all
cells bordering cells sampled in the previous step that contained sign and continuing to sample
in this manner until the cluster is surrounded by vacant cells (Thompson 1992; Thompson
and Seber 1996).
Observability is a source of variation in presence-absence surveys just as in surveys directed
at abundance, because not all cells or sample plots containing tiger sign will be identified
(sign can be “missed”). Tiger “presence” is assumed to be determined without error, such that
detections of tiger sign are not confused with sign of other large cats and vice versa. Tiger
“absence” can reflect either true absence, or presence with non-detection. Different possible
approaches exist for estimating detection probability, which we define as p = probability that
tiger sign is detected, given tiger activity (hence sign presence) in the cell or sample plot.
Estimating Observability
Multiple observers
One approach involves multiple (two or more) independent observers. The observers survey
s cells or sample plots independently (e.g., on different consecutive days, or perhaps on different
2-hour periods within the same day), and the objective is to estimate the number of these cells
that contained sign (denote this number as r).
Assume, for this example, that two observers are used. Then the statistics (the actual data)
that result from their efforts to detect sign in each cell are:
x10 = number of the s cells in which only observer one detects sign,
x01 = number of the s cells in which only observer two detects sign,
x11 = number of the s cells in which both observers detect sign.
Detection probabilities for observers one and two can be estimated as:
x11
pˆ 1 =
x11 + x 01
(4.2)
x11
pˆ 2 =
x11 + x 10
These estimators should correspond with intuition. For example in order to estimate p1, we
simply sum the cells at which we know sign was present because observer two detected it
( x11 + x01 ) , and compute the proportion of these cells at which observer 1 detected sign. The
estimated overall (both observers) detection probability is given by:
pˆ = 1 − (1 − pˆ 1 ) (1 − pˆ 2 ) (4.3)
The above expression simply notes that both observers must fail to detect sign in order for the
cell or sample plot to be recorded as a nondetection. The estimated number ( rˆ ) of the s cells
that contained sign is then given by:
x11 + x10 x + x01 x + x10 + x01 ( x11 + x10 ) ( x11 + x0
rˆ = = 11 = 11 = (4.4)
pˆ 1 pˆ 2 pˆ x11
The above is simply the Lincoln-Petersen estimator (e.g., Seber 1982). In this case, the estimator
corresponds to the number of occupied cells (cells containing sign). The estimator is based on
the principles discussed in Chapter 3 and specifically on the canonical estimator of (3.2). For
example, in the first equality of (4.4), we just take the number of cells at which sign was
detected by observer one and then divide that count statistic by the estimated detection
probability for observer one. Similarly, we can use the count of observer two divided by the
estimated detection probability of that observer, or the total count of both observers divided
by their estimated detection probability. These three different ways of writing the canonical
estimator for this particular sampling situation all lead to the same estimator (i.e., the same
function of statistics that arise from the field sampling). However, in practice it is best to use
Chapman’s (1951) bias-adjusted estimator (Seber 1982), as the estimator in (4.4) is slightly
biased when sample sizes are small. See Chapter 11 and equations 11.5 and 11.6 for the
Chapman estimator and its associated variance.
Although the number of occupied cells can be estimated with 2 investigators as described
above, there are advantages in both precision and estimator robustness in using multiple
investigators. By “precision” we refer to the variance or the 95% confidence interval of the

30 31
 Statistical Concepts: Assessing Spatial Distributions
estimator, with a precise estimate having a small variance and a narrow confidence interval.
By “robustness” we refer to the ability of the estimator to produce estimates with relatively
small bias even when underlying assumptions are not met. If five or more investigators can be
sent to each cell or sample plot, then the set of closed capture-recapture models described by
Otis et al. (1978; see Chapter 11) can be used. Model selection procedures implemented in the
software recommended for these models, CAPTURE (Rexstad and Burnham 1991), help the
user select the model most likely to be useful with the data set. The different models make
different sets of assumptions about sources of variation in capture or detection probabilities,
and the model selection algorithm of CAPTURE directs the user to the set of assumptions,
and hence to the model and estimator, that appear to be appropriate for each data set (see
Chapter 11).
Multiple visits by one observer
Still another approach to estimating detectability in presence-absence surveys requires
multiple (e.g., two) visits by a single observer. The approach to estimation is that used in
estimation with a removal model (Seber 1982). The observer initially surveys s cells (denote
these initial visits as sampling occasion one), and returns at a later time (denote as sampling
occasion two) to resurvey cells at which no sign was found on occasion one. Again, we want
to estimate the number of the s cells that contained sign (denote as r). The following statistics
are obtained:
x1 = number of cells at which sign was found on occasion one,
x2 = number of resurveyed cells (no sign detected at occasion one) at which sign was
found on occasion two.
In the usual case where x1>x2, the estimator for the number (rˆ ) of the s cells that contained
sign using this approach is (e.g., Seber 1982:318)
x
2
rˆ = 1 (4.5)
x1 − x2
with variance estimator:
x x ( x1 + x2 )
2 2
vaˆr(rˆ) ≈ 1 2
(4.6)
( x1 − x 2 ) 4
The above is known as the Zippin removal estimator for the special case of two samples (see
Otis et al. 1978). The estimated overall (both visits) detection probability is given by:
32
Statistical Concepts: Assessing Spatial Distributions
x1 + x 2 x
pˆ = = 1 − ( 2 )2 (4.7)
rˆ x1
This estimator for detection probability corresponds both to intuition and the conceptual
framework presented in Chapter 3, in that we simply divide the number of cells at which sign
was detected, by the estimated number of cells at which sign was present (occupied cells).
The two-visit approach described above can be extended to >2 visits as well. In this case, for
example, cells at which no sign is found on visits one and two are revisited a third time, etc.
Models Mb and Mbh of Otis et al. (1978) are the appropriate models in this case, and program
CAPTURE (Rexstad and Burnham 1991) can be used to compute estimates (see Chapter
11). More visits typically lead to greater precision and permit estimation under a model in
which occupied cells can differ with respect to the ease of finding sign.
Estimation Under Different Sampling Designs
Simple random sampling
Regardless of whether the number of occupied cells or sample plots, r, is estimated using a
removal estimator (single investigator) or a capture-recapture estimator (multiple observers),
recall that the objective is to estimate the proportion of area (cells) occupied by tigers. We
now have an estimate of the number of sampled cells that are occupied by tigers, and we
know the number of cells that were sampled, (i.e., the number of cells that were searched for
sign). Under simple random sampling (i.e., assuming that the surveyed cells were selected at
random from all possible cells in the area of interest), we can simply estimate the proportion
of interest as:
rˆ
ψˆ = (4.8)
s
This estimator is a very natural one, as we are simply estimating the proportion of cells that
are occupied by dividing the estimated number occupied by the total cells sampled. The
variance for the estimated proportion of area occupied can be estimated as:
vaˆr( rˆ) ψˆ (1 − ψˆ )
vaˆr(ψˆ ) ≈ + (4.9)
s2 s
where vaˆr(rˆ) is estimated as specified for the selected method for estimating r. For example,
if we used two visits by a single observer, then we would compute vaˆr(rˆ) using equation 4.6.
Example: As an example, consider a 2000 km2 area of interest that has little history of tiger
33
 Statistical Concepts: Assessing Spatial Distributions Statistical Concepts: Assessing Spatial Distributions

research or investigation, and assume that there is interest in estimating the fraction of the estimate is divided by the fraction of the total cells in the stratum that were sampled, aj, in
area that is occupied by tigers. Further assume that the area has been subdivided (e.g., using order to estimate the number of cells containing sign, Rˆ j , in each stratum. These stratum–
GIS- see Chapter 6) into a grid of 10 km2 cells or sample units. Thus, there are 200 of these specific estimates of occupied cells are summed to yield Rˆ :
cells in the entire area of interest. Assume that the question of proportional occupancy has
been operationally defined as: “What proportion of the cells contain evidence of tiger activity?” rˆj
Rˆ = Σ = Σ Rˆ j (4.10)
Assume that 30 plots have been randomly selected from the sample population of 200 plots j αj j
for estimating the proportion of the plots that are occupied by tigers. Finally, assume that a
single investigator locates each selected plot, travels to it, and spends two days searching for ψ for the entire area is then estimated by dividing the estimated number of occupied
sign of tiger activity (pugmarks, scats, etc.). Plots on which no tiger sign is found are then cells in the entire area by the total cells:
revisited, and the same investigator conducts an additional two day search. These data are
suitable for use with the removal model approach for estimation of the number, and proportion, Rˆ
ψˆ = (4.11)
of plots with tiger activity. Using the removal model notation defined above, assume that this S
investigation yields x1=13 plots or cells on which sign of activity was found on the initial visit
and search of the cell. Further assume that sign was found on the second visit for x2=4 of the Example: As a simple illustration of stratified random sampling, assume that a 4000 km2
17 cells that were revisited. Using these values in equation (4.5) yields the following estimate area of interest is divided into a Sal forest stratum (denote as stratum 1, containing 1000 km 2)
of the number of cells or plots with tiger activity: and a riparian grassland stratum (denote as stratum 2, containing 3000 km2) and that both
strata have been subdivided into 10-km2 cells for the purpose of sampling. Assume that the
(13) 2 two-observer approach has been used with a sample of 20 cells in stratum 1 (thus
rˆ = ≈ 19
(13 − 4) 20
α1 = = 0.20) to yield an estimate of rˆ1 = 8 cells occupied by tigers in stratum 1. Further
100
with associated variance (from equation 4.6)
40
assume that 40 cells in stratum 2 were sampled (so α 2 = ≈ 0.133 ) to yield an estimate of
300
(13) 2 ( 4) 2 (13 + 4)
vaˆr(rˆ) ≈ ≈ 7 .0 rˆ2 = 26 cells. Application of equation (4.10) yields:
(13 − 4) 4
8 26
The estimated proportion of cells occupied (equation 4.8) is then given by Rˆ = Rˆ1 + Rˆ 2 = + ≈ 235
0.200 0.133
rˆ 19
ψˆ = = ≈ 0.63 From this estimated number of occupied cells in the entire area, the proportion of area occupied
s 30 can be estimated using equation (4.11) as:
with estimated variance (equation 4.9)
235
ψˆ ≈ ≈ 0.59
7.0 0.63(1 − 0.63) 400
vaˆr(ψˆ ) ≈ 2
+ ≈ 0.0078 + 0.0078 = 0.0156
(30) 30 So the estimated proportion of area occupied in the entire area is 0.59. It is also possible to
estimate the proportion of area occupied for each stratum using equation (4.8) for each stratum:
Stratified random sampling
rˆ1 8
Under stratified random sampling (we randomly select cells from within each stratum), we ψˆ 1 = = = 0.40
s1 20
use the estimated detection probability to estimate (e.g., using equations 4.4 or 4.5) the total
number of sampled cells that are occupied in each stratum, rˆj , where j indexes stratum. This

34 35
 Statistical Concepts: Assessing Spatial Distributions
rˆ2 26
ψˆ 2 = = = 0.65
s 2 40
The estimated proportion of area occupied for the entire area can also be computed as a
weighted average of the stratum-specific estimates, where the weights are based on the
proportional representation of the stratum in the entire area. Stratum 1 comprised 100/400 =
0.25 of the area, whereas stratum 2 comprised 300/400=0.75 of the entire area. The weighted
average of the stratum-specific estimates of proportion occupied is computed as:
ψˆ = 0.25ψˆ 1 + 0.75ψˆ 2 = (0.25)(0.40) + (0.75)(0.65) ≈ 0.59
Thus, the two approaches yield identical estimates of ψ .
Adaptive sampling:
The situation is more complicated under adaptive sampling, but an estimator for R is developed
using the pˆ and the cluster-specific locations of cells at which sign was found. The resulting
Rˆ is then used in expression (4.1) to estimate ψ . The reader is referred to Thompson (1992)
and Thompson and Seber (1996) for details of adaptive sampling methods.
MAPPING THE SPATIAL DISTRIBUTION
Exercises directed at mapping the spatial distribution of tigers or their prey generally ignore
the issue of detection probabilities and assume that animals are detected with probability 1
(so if animals are present in an area, then they are always detected). Using the methods
described above, it is possible to estimate the number of occupied cells that are “missed”
during sampling, but it is not possible to specify the exact location of missed cells. Thus, the
primary design consideration for mapping the entire distribution of tiger or prey animals will
likely be spatial sampling. We offer a few conjectures about survey designs for the purpose of
mapping overall tiger or prey distribution.
If there is no strong clustering of tigers, then a systematic or uniform sampling pattern may
be best. Our rationale is that systematic, uniform sampling can minimize the average distance
of unsurveyed cells to surveyed cells. In general, the farther an unsurveyed cell lies from
surveyed cells, the more difficult it is to make inference about its possible occupancy. If
strong clustering does exist, then adaptive cluster sampling (Thompson 1992; Thompson and
Seber 1996) may be worthy of consideration.
In either case, it may be possible to use methods based on the spatial correlation structure of
36
Statistical Concepts: Assessing Spatial Distributions
cell occupancy to draw inference about unsurveyed cells (“Kriging” is a method based on this
idea, Thompson 1992). Another possibility is to use the spatial pattern of covariation of tiger
presence with some covariate whose distribution is better known (e.g., it can be easily measured)
and to model tiger presence as a function of the covariate.
Finally, note that mapping may involve characteristics other than presence-absence. For
example, it might be possible to design special surveys for mapping areas with tiger cub
production for use in identifying population source and sink areas.
Issues relating to mapping the distribution of animals are covered more thoroughly in Chapters
5 and 6. We believe that such mapping exercises can be most useful when they are conducted
in conjunction with an appropriate spatial sampling design. Under this approach, all locations
need not be sampled, but the critical point is that all locations have known a priori probabilities
of being included in the sample. This condition permits reasonable inference when only a
subset of potential locations is surveyed.
We also believe that it may be useful to consider mapping exercises that incorporate the issue
of detectability. For example, it would be possible to develop maps with different shadings or
contour lines marking areas with different rates of occupancy, as estimated using the methods
described above for estimating proportion of area occupied. Such maps would provide an
accurate picture of animal distribution and would not be based on an incorrect assumption
that all animals present in surveyed areas are detected.
LOOKING AHEAD
The methods presented above for estimating the proportion of area occupied by a species are
relatively new, at least in one sense. The estimators are borrowed from capture-recapture
modeling and were developed decades ago, but their application to data arising from repeat
surveys for animal presence is new. These methods are perceived as having good potential
and are thus the subject of ongoing research. For example, software has been recently developed
for the purpose of modeling detection probability and the probability of occupancy from
presence-absence survey data (Mackenzie et al. 2002). This software is based on extensions
of the methods presented in this chapter and permits estimation of both detection probability
and occupancy as functions of covariates. Such modeling would permit formal tests of ideas
about animal preferences for certain types of habitat, etc.
In addition, we are now working on development of models that use data from presence-
absence surveys conducted over multiple years. The models view changes in occupancy status
37
 Statistical Concepts: Assessing Spatial Distributions CHAPTER 5

as functions of local extinction and colonization probabilities, and permit direct estimation of FIELD SURVEYS: ASSESSING SPATIAL DISTRIBUTIONS OF
rates of local extinction, colonization and change in occupancy (i.e., increases or decreases in TIGERS AND PREY
the proportion of area occupied over time). Such models could be used in conjunction with
large-scale presence-absence surveys to draw inferences about temporal changes in population
status. Such models and surveys seem well-suited to large-scale monitoring. K. Ullas Karanth1, N. Samba Kumar2 and Raghunandan S. Chundawat3
Wildlife Conservation Society (International Programs), Bronx, New York, NY-10460-1099, USA.
1

Email: <ukaranth@wcs.org>
REFERENCES
Centre for Wildlife Studies, 26-2 Aga Abbas Ali road (Apt. 403), Bangalore, Karnataka-560 042, India.
2

Email: <wcsind@bgl.vsnl.net.in>
Chapman, D.G. 1951. Some properties of the hypergeometric distribution with application to
zoological censuses. University of California Publications in Statistics 1:131-160. Wildlife Institute of India, P.B. 18, Chandrabani, Dehradun, Uttaranchal-248 001, India.
3

Email: <pnptiger@mantraonline.com>

Mackenzie, D.I., Nichols, J.D., Lachman, G.B., Droege, S., Royle, J.A. and Langtimm,
C.A. 2002. Estimating site occupancy rates when detection probabilities are less
than one. Ecology (in press).
INTRODUCTION

Conservation and management planning require basic information on the spatial distribution
Otis, D.L., Burnham, K.P., White, G.C. and Anderson, D.R. 1978. Statistical inference from
of tigers and prey species at regional and national levels, at an appropriate scale (Karanth
capture data on closed animal populations. Wildlife Monographs 62:1-135.
and Nichols 2000). Therefore, the most useful activity that can be taken up at regional or
countrywide levels is the assessment and monitoring of the spatial distribution of tiger (and
Rexstad, E. and Burnham, K.P. 1991. User’s guide for interactive program CAPTURE.
prey) populations using field surveys (Chapter 1). The statistical concepts underlying such
Abundance estimation of closed animal populations. Colorado State University,
surveys of spatial distribution were covered in Chapter 4, under the general estimation and
Fort Collins, CO, USA.
sampling framework described in Chapter 3.
Seber, G.A.F. 1982. The estimation of animal abundance and related parameters. MacMillan,
As we saw in Chapter 1, given the critical status of tigers and the substantial investments
New York, NY, USA.
being made in tiger conservation, wildlife managers and conservation agencies primarily
need clear and reliable answers to the following basic questions regarding distribution of
Thompson, S.K. 1992. Sampling. Wiley, New York, NY, USA.
tigers:
Thompson, S.K. and Seber, G.A.F. 1996. Adaptive sampling. Wiley, New York, NY, USA.
1. What is the distributional range occupied by different individual tiger populations in the
country or region?

2. Where are individual tiger populations increasing their ranges and where are the ranges
fragmenting and shrinking?

3. Within the entire tiger range, what is the proportion of the area occupied by breeding tiger
populations?

To address these questions, it is necessary to establish valid protocols for sampling, recording
and mapping distributions of tigers and their prey species over large regions (Karanth 1999;

38 39
 Field Surveys: Assessing Spatial Distributions of Tigers and Prey
Karanth and Nichols 2000). Such sample surveys of distribution can provide information of
management and scientific relevance, even under field conditions characterized by scarcity of
resources and lack of trained manpower that prevail in most parts of the tiger range.
Sample surveys of distribution can provide us with an accurate picture of metapopulation
structures of tigers over large regions (Smith et al. 1999). They help us identify potential
habitat corridors, landscape connectivity, and dispersal routes used by transient tigers. When
such surveys are conducted periodically (e.g., each year) as part of a monitoring program,
then resulting data can be used to assess the influence of landscape characteristics on
metapopulation vital rates (local extinction and colonization rates) and changes in occupancy.
Sometimes, these surveys can help us identify and deal with prevailing or emerging threats to
tigers, prey and habitats.
Although it is desirable to carry out field surveys over the entire tiger distribution range, for
various reasons, it may not be possible to do this every year. In such cases, some useful
information on spatial distribution over a wider area can also be gathered from questionnaire
surveys of knowledgeable informants. Even if field surveys are carried out, it may be useful
to conduct additional questionnaire surveys to supplement the information. Information about
emerging threats such as poaching, adverse impacts on habitats and the effectiveness of
protective measures also can be effectively generated from questionnaire surveys.
In the following two sections, we explain the survey protocols that we believe will lead to
reliable and cost-effective assessments of spatial distributions of tigers and prey species over
large regions. These protocols may need some modifications to suit local conditions.
FIELD SURVEYS
Recording Signs
Tigers are secretive, wide-ranging animals, distributed over an extensive area. For example,
the extent of potential tiger habitat in tropical Asia is estimated to be over 1,200,000 square
kilometers (Wikramanayake et al. 1999). They are difficult to observe in the field and therefore
signs like tracks, scats and kills are important means for assessing the status of tiger populations
over large regions.
Because the viability of tiger populations is chiefly a function of prey abundance (Karanth
and Stith 1999; Miquelle et al. 1999), questions about tiger status can be answered better if
tiger surveys are carried out simultaneously with surveys of the principal prey species. Surveys
of distribution of the tiger’s prey are generally easier to carry out, because ungulates are
40
Field Surveys: Assessing Spatial Distributions of Tigers and Prey
relatively more abundant, easier to detect and generate much more tracks and dung than
tigers themselves. In this chapter, we refer to all such indirect evidences as animal sign.
Field sample surveys of sign should record such information as tracks of adult tigers and
cubs, kills, scats and sightings of prey or detection of prey sign. Such surveys may typically
require 3 to 15 days, and should be carried out at least once a year. Region-wide surveys
necessarily require a large number of personnel like government staff, laborers or volunteers.
Recognizing Tiger and Prey Signs
Personnel participating in the field survey should be able to correctly recognize the signs of
tigers and prey species that they encounter in the field. Most often tigers are not seen, but
their signs are. Although in some areas prey species are encountered during field surveys,
quite often even their distribution has to be assessed from signs.
Usually, experienced survey personnel can distinguish tiger tracks from leopard tracks. Tracks
that look like leopard tracks, but accompany a definite tiger track, often indicate a tigress
moving with her cubs. Survey personnel should also be able to recognize the principal prey
species and their signs and calls. They should have a thorough geographical knowledge of the
area they are going to survey.
Several publications describe how to recognize and record tiger and prey signs in the field
(van Strien 1983; Rabinowitz 1997; WWF-Nepal 1998; McDougal 1999). In this manual we
have provided illustrations of common large carnivores and prey species of tigers that occur
in tropical Asia and of the tracks they leave (Appendix 1.1 and 1.2). These materials can be
used to familiarize survey personnel with the survey objects. However, we reiterate that
manuals are not a substitute for actually looking at animals or signs and gaining practical
experience in accurate recognition. We emphasize that it is absolutely essential that collections
of animal specimens (or at least their skins), and specimens of tracks, scats and dung must be
built up in each reserve. Such specimens should be used for training field survey personnel.
Detecting Tiger and Prey Sign in the Field
Tracks of tigers and leopards are readily detected wherever substrate conditions are appropriate.
In the tropics, such conditions are met on dusty roads, sandy riverbeds, riparian deltas and
alluvial soils. Unfortunately, in most areas of the tropics, tiger tracks are difficult to find
because of unsuitable soil, rainfall and leaf-fall. Where tiger tracks are not easily detected,
sometimes tiger scats can be used for range-mapping purposes, although usually fewer scats
than tracks are encountered (Figure 5.1). Similarly, surveys must also record the signs of
principal prey species (including livestock). Sightings of prey species, tracks, dung, calls are
the type of data gathered from field surveys.
41
 Field Surveys: Assessing Spatial Distributions of Tigers and Prey Field Surveys: Assessing Spatial Distributions of Tigers and Prey

Manpower Resources for Field Surveys

An adequate number of personnel should be assigned to survey the area being assessed (e.g.,
a reserve, park, sanctuary, forest range, reserved forest as the case may be). Personnel must
have a full day (morning to evening) to complete the search in each sample unit of the field
survey.

Remember that you are not trying to count every individual tiger to get a total count (census)
for the surveyed area. Therefore, surveys need not be conducted in neighboring areas

Figure 5.1 - Field survey for tiger signs in progress (Inset: Basic tools for field surveys).
simultaneously. This flexibility increases your ability to deploy survey personnel optimally.
Your aim should be to cover the survey area once in a year, during the same month/season,
within a short time span (3-15 days), in a standardized manner to make the detection
probabilities similar, to the extent possible.

Sometimes, survey personnel who possess adequate field skills may lack the writing skills to
systematically record the data. Therefore, at least one person who can record the data should
be on each survey team.

Often, large numbers of staff or local volunteers can be deployed over short periods of time.
Such large manpower resources are necessary for simultaneous landscape-scale surveys of
tiger and prey distribution. Alternatively, smaller teams of trained surveyors can sequentially
cover the areas to be surveyed, extending the survey over longer periods but reducing the
requirement of personnel. For example, let us say that tiger sign surveys over a large region
of several hundred square kilometers of forest can be completed in a day by 100 teams of
three persons each. On the other hand, if there are only 10 such teams, the same work can still
be completed, but, over10 days. Your choice here depends on a compromise between the
quality and numbers of personnel, and the time available to you to complete the survey.

The large manpower needed for region-wide surveys of tiger-prey distribution can be met
9

from a variety of sources: Staff of wildlife or forestry departments, other government staff,
tour guides, local naturalists or volunteers from non-governmental organizations.
6

Material Resources
8
4

It is necessary to have reference collections of plaster casts of tracks and specimens of scats,
2

8. Aluminium Strip

dung, pellets of animals being surveyed: Tigers, leopards, other large carnivores and principal
6. Measuring Tape
3. Pugmark Tracer
1. Plaster of Paris

7. Water Bottle

prey species. Such reference collections will have to be built up from specimens of known
5. Data Form

origin obtained either from captivity or in the wild as and when opportunities arise. These
7

reference collections will be helpful in training the field staff to identify signs that they observed
4. Pens

9. Mug
2. Bag
1

5
3

during surveys in the field (Appendices 1.1 and 1.2).

42 43
 Field Surveys: Assessing Spatial Distributions of Tigers and Prey
The basic survey equipment and consumable materials needed for field surveys (Figure 5.1
[Inset]) are listed below:
1. Plaster of Paris and associated equipment for collecting track casts
2. Polythene or paper bags to collect scats
3. Data recording forms
4. Adequate number of 1: 50,000 or at least 1: 250,000 scale, georeferenced maps.
5. Measuring instruments – tapes, calipers etc.
6. Binoculars, Cameras and GPS units, if available.
Field Survey Protocols
Choose the best season and month of the year for the survey, based on your knowledge of
field conditions: For example, soon after first showers or snowfall, when there is less leaf
litter, when survey personnel are free of other tasks or when volunteers are easier to find. The
whole point is to do a good survey when personnel are available, logistics are tractable and
animal signs easier to find. After you have chosen the best time for the survey, try to conduct
the survey in roughly the same period every year thereafter. Such standardization is helpful
for monitoring effort (Chapter 7). The best time for field surveys is likely to vary from region
to region even within a country.
As noted in Chapters 3 and 4, well-designed field surveys must include consideration of both
spatial sampling and detectability. Spatial sampling will involve subdividing the total area of
interest (e.g., a park or a large region) into a number of potential sampling units. The size of
the unit will be dictated by the amount of area that a survey team can search during a single
period (e.g., one day). Under simple random sampling some number of these potential sample
units is randomly selected to be surveyed on the ground. Under stratified random sampling,
the entire area is subdivided into two or more strata or areas among which tiger abundance is
expected to vary (e.g., areas with low, medium and high prey densities). Then some number
of sample units is randomly selected from each stratum. Spatial sampling design simply
involves this selection of sample units to be actually surveyed by ground crews.
For the selected sample units, detectability can then be estimated using either the multiple
observer approach or the approach of a single observer and multiple visits as discussed in
Chapter 4. Under the multiple observer approach, two different survey teams are sent to the
same sample unit(s) on different days. Time separating the two visits should be short, but
long enough that human sign left by the first survey team does not lead the second team to
44
Field Surveys: Assessing Spatial Distributions of Tigers and Prey
animal sign. Under the single observer approach, a single survey team revisits for a second
time sample units on which no sign was detected the first time. Under both multiple and single
observer approaches, the data resulting from survey efforts is a list of species that were
detected or not detected on each survey visit. These presence-absence data collected on the
repeat visits then form the basis for estimating detectability, and hence the total proportion of
sample units that is occupied (Chapter 4).
Train your survey personnel well to recognize and record signs of tigers and prey animals.
Identify and designate field survey teams of 3-4 persons (or whatever number local conditions
dictate), including in each team at least one member capable of reliably recognizing animals
and their signs. At least one member of the team must be capable of writing down the
information and later transferring it to the maps.
Determine the number of days of actual fieldwork possible after carefully considering the
number of competent survey teams available and the logistical problems involved (rivers to
be crossed, topography and terrain problems, vehicles, accessibility, safety of personnel,
snow, rainfall, holidays etc.). It is very important to be realistic and practical about the areas
to be covered, distances to be walked, and how and by whom the survey will be conducted
and supervised.
If for any reason it is not possible to cover an area either fully (or even partially) please report
this fact unambiguously. Such partial surveys could result from natural calamities, social
unrest, shortage of personnel, or any number of other local factors. If the fact that the survey
was not carried out or was abandoned is reported correctly, then it is not a problem to deal
with the results during the subsequent analyses. If such facts are not reported the survey data
will be vitiated beyond repair.
Please recognize clearly that the entire area cannot be surveyed in most cases (the spatial
sampling issue; Chapters 3 and 4): Only a part of the total area will be surveyed, even in well-
executed, intensive surveys. Also recall, the survey teams will not detect all animal sign
present even within the areas they are able to visit (Observability problem: Chapters 3 and 4).
Don’t be daunted by these problems: Remember that in a properly conducted sample survey,
the protocols described in this manual will give you the power to make inferences about un-
surveyed areas and undetected animal signs. Therefore, accurately record the key piece of
information, sampling effort, in terms of distances covered, localities and timings as described
in the model survey data forms provided in this manual (Appendix 1.3).
If voucher specimens of tiger or leopard tracks are required, obtain those in the
form of plaster casts, photographs or tracings. However, please note that we are not
45
 Field Surveys: Assessing Spatial Distributions of Tigers and Prey
trying to identify individual tigers or leopards, only ascertaining presence or absence of
these species.
Even in the best of tiger habitats it is not always easy to find tiger sign on the day you do the
survey. Therefore, even after a diligent search, some field survey teams may not find any tiger
signs at all. Please recognize that such outcomes are not unusual and they are likely to increase
at lower tiger and prey densities. In most habitats outside well-protected reserves, tiger and
prey densities will be very low and their sign very scarce. Such low- density areas now cover
more than 90% of the tiger distribution range (Karanth 2001). In most parts of the tiger
range, absence of suitable soil, leaf-litter and grass render it difficult to find tiger and prey
sign. In some cases, continuous rainfall, tidal changes, and animal activity may obliterate
signs.
Not finding tiger or prey sign does not necessarily mean that the survey team is not doing its
job. It also does not necessarily mean that tigers and prey are absent from the area. It is often
a matter of chance that sign is detected (Chapter 4). It should be emphasized to survey
personnel that the utility of their work is not judged by whether or not they detect sign, but by
whether they follow survey guidelines and instructions.
Even when no signs are found, well-executed sample surveys are useful in making inferences
about probabilities of detecting sign in the surveyed region. Therefore, do not admonish
survey teams that fail to report signs, as long as they have carried out the survey as directed.
Compelling survey teams to come up with evidence, or denigrating them if they do not find
tiger sign, may force them to fake the data in a manner that is practically undetectable. This
may lead survey personnel to cheat: Reuse old plaster casts, lift multiple prints of the same
set of tracks, inflate the number of prey animals seen or simply cook up the field data. Such
vitiated data generated under pressure are of no value whatsoever, and will undermine the
good work done by other survey teams. Bad data will lead you to false conclusions: In such
an event you are much better off without any data.
It is very important to georeference the field survey data. What this means is that the data on
tiger sign, specimens collected, and other records, should be linked to a map that has accurate
latitudes and longitudes (or UTM coordinates) marked on it. A 1:50,000 or at least 1: 250,000
scale map should be used to record the locations of field observations. Preferably all location
data should be recorded at an accuracy of 100 meters, or at least to within 1000 meters on the
map. If good maps are available, this task becomes easier. It is crucial to transfer the information
from the surveys to georeferenced maps without delay.
Most importantly, the survey teams must record the sampling effort accurately in terms of
46
Field Surveys: Assessing Spatial Distributions of Tigers and Prey
number of persons per team, number of teams sent out, time spent surveying and distance or
area covered by each survey team etc., as indicated in the model data form (Appendix 1.3).
These are all quantitative measures of sampling effort.
Each team’s work on each different date should be treated as a separate survey, given a
unique serial number and recorded on a separate form. Data forms must be numbered and
cross-linked to the specific georeferenced maps. For example, a particular data form may be
numbered 22 in Nagarahole Reserve and linked to a map titled Karnataka State -18 in India.
A particular specimen collected during this particular survey (say Tiger Scat-4) should be
cross-linked to both the related data form and the map. Sufficient copies of data forms should
be available before the survey is commenced.
QUESTIONNAIRE SURVEYS
Questionnaire surveys gain importance if large regions cannot be surveyed using field teams
as described earlier. Interviews of local informants such as hunters, local naturalists and park
personnel can be useful, if they are truly knowledgeable: Informants should not be randomly
drawn individuals from the local population!
Often, knowledgeable informants may not be literate to fill out the data forms and mark
locations on maps: They may have to be assisted by others. Questionnaire surveys should be
conducted by trained personnel capable of assessing the quality of information provided by
informants (Figure 5.2). If an informant appears to be ignorant or untruthful, the data should
be discarded ruthlessly. If informants and survey personnel of adequate caliber are not found,
there is no point in carrying out questionnaire surveys.
The results of questioning each individual informant must be treated as a distinct sample.
Each such sample of data must be recorded on a fresh questionnaire survey form. As in the
case of field surveys, the forms must be systematically numbered and cross-linked to
georeferenced maps.
Although ideally contour maps should be used, in their absence, other maps such as Reserve
Maps, Forestry Maps, District Maps etc., can also be used. However, It is important that all
maps on which survey information is recorded are properly georeferenced: They should have
a scale, identify prominent geographic features and display latitudes and longitudes accurately.
A specimen questionnaire form is shown in Appendix 1.4. Please note that not all the information
sought in this particular specimen is related to spatial mapping of tigers and prey. This form
can be modified to suit your local needs.
47
 Field Surveys: Assessing Spatial Distributions of Tigers and Prey
Figure 5.2 - Questionnaire survey for tiger presence in progress.
ORGANIZING THE SURVEY DATA FOR MAPPING AND ANALYSIS
The data from the field surveys or questionnaires described above will be in the form of
several maps and data forms linked to them. The investigator should examine all these forms
and maps to correct obvious errors, remove ambiguities, discard questionable data and try to
fill in any missing information by interviewing the survey team. This must be done immediately
after the surveys are completed. The investigator may physically re-check a certain proportion
of the data by personally visiting the area and verifying the evidence. Such random checks
enhance the integrity of data collection.
When the survey is completed, the investigator must ensure that the resulting data forms and
maps are intelligible to the person preparing the spatial distribution maps or doing other
analyses (Chapters 4, 6 and 7).
48
Field Surveys: Assessing Spatial Distributions of Tigers and Prey
REFERENCES
Karanth, K.U. 1999. Counting tigers, with confidence. Pages 350-353 in Riding the tiger:
Tiger conservation in human-dominated landscapes (Eds: J. Seidensticker, S. Christie
and P. Jackson). Cambridge University Press, Cambridge, UK.
Karanth, K.U. 2001. The way of the tiger: Natural history and conservation of the endangered
big cat. Voyageur Press. Stillwater, MN, USA.
Karanth, K.U. and Nichols, J.D. 2000. Ecological status and conservation of tigers in India.
Final Technical Report to the US Fish and Wildlife Service (Division of International
Conservation), Washington, DC, and Wildlife Conservation Society, New York. Centre
for Wildlife Studies, Bangalore, India.
Karanth, K.U. and Stith, B.M. 1999. Prey depletion as a critical determinant of tiger population
viability. Pages 100-113 in Riding the tiger: Tiger conservation in human-dominated
landscapes (Eds: J. Seidensticker, S. Christie and P. Jackson). Cambridge University
Press, Cambridge, UK.
McDougal, C. 1999. You can tell some tigers by their tracks with confidence. Pages 190-191
in Riding the tiger: Tiger conservation in human-dominated landscapes (Eds:
J. Seidensticker, S. Christie and P. Jackson). Cambridge University Press, Cambridge,
UK.
Miquelle, D.G., Smirnov, E.N., Merrill, T.W., Myslenkov, A.E., Quigley, H.B., Hornocker,
M.G. and Schleyer B. 1999. Hierarchical spatial analysis of Amur tiger relationships
to habitat and prey. Pages 71-99 in Riding the Tiger: Tiger conservation in human
dominated landscapes (Eds: J. Seidensticker, S. Christie and P. Jackson), Cambridge
University Press, Cambridge, UK.
Rabinowitz, A. 1997. Wildlife field research and conservation training manual. Wildlife
Conservation Society, New York, NY, USA.
Smith, J.L.D., McDougal, C., Ahearn, S.C., Joshi, A. and Conforti, K. 1999. Metapopulation
structure of tigers in Nepal. Pages 176-189 in Riding the tiger: Tiger conservation in
human-dominated landscapes (Eds: J. Seidensticker, S. Christie and P. Jackson).
Cambridge University Press, Cambridge, UK.
van Strien, N.J. 1983. A guide to the tracks of mammals of W. Indonesia. Unpublished.
49
 Field Surveys: Assessing Spatial Distributions of Tigers and Prey CHAPTER 6

Wikramanayake E.D., Dinerstein, E., Robinson, J.G., Karanth, K.U., Rabinowitz, A., Olson, SPATIAL DISTRIBUTIONS OF TIGERS AND PREY: MAPPING
D., Matthew, T., Hedao, P., Conner, M., Hemley G. and Bolze D. 1999. Where can AND THE USE OF GIS
tigers live in the future? A framework for identifying high priority areas for the
conservation of tigers in the wild. Pages 255-272 in Riding the tiger: Tiger
conservation in human-dominated landscapes (Eds: J. Seidensticker, S. Christie Bradley M. Stith1 and N. Samba Kumar2
and P. Jackson). Cambridge University Press, Cambridge, UK.
United States Geological Survey, Florida Caribbean Science Center, Sirenia Project. 412 NE 16th Ave;
1

Rm 250, Gainesville, FL-32601, USA.

WWF-Nepal 1998. Tiger Manual: Indirect field study techniques for the kingdom of Nepal. Email: <stith@isof.net>
World Wildlife Fund-Nepal, Kathmandu, Nepal.
Centre for Wildlife Studies, 26-2 Aga Abbas Ali road (Apt. 403), Bangalore, Karnataka-560 042, India.
2

Email: <wcsind@bgl.vsnl.net.in>

INTRODUCTION

In Chapter 5, we examined how surveys of the presence, absence and distribution of tigers
and their prey species are carried out in the field. A fundamental objective of such field
surveys is to map the spatial distribution of tigers and their prey species, and the various
human factors that affect both tigers and prey. In this chapter, we describe how to achieve this
objective by collating and summarizing the information from field survey forms to produce
maps showing the distribution of tigers, prey species, and other related variables. In the past,
this process of collating information and producing maps was done laboriously by hand.
Now, the computer technology known as Geographic Information Systems (GIS) provides a
powerful tool that greatly facilitates this process. Below, we provide a simple overview of
GIS techniques, emphasizing the steps needed to map and analyze distributions of tigers and
their prey. A detailed treatment of the use and application of GIS is beyond the scope of this
manual; such information is available from the following references: Star and Estes 1990;
Koeln et al. 1994; Burrough and McDonnell 1998; Morrison et al 1998; Corsi et al. 2000.

DATA SOURCES

For the purpose of mapping tiger and prey distributions, we rely on two fundamentally different
sources of data. The first source is the field and questionnaire survey data, and the second is
ancillary data obtained from external sources. Detailed protocols for collecting the field and
questionnaire survey data have been described earlier (Chapter 5). The ancillary data can be
almost any kind of appropriately scaled data, including satellite imagery, forest cover maps,
topographic sheets, government statistics, etc. GIS provides a powerful tool for integrating
and synthesizing the field survey data with disparate types of ancillary data, and ensuring
that these data will have the proper spatial relationships to one another and will display
properly when combined on individual maps.

50 51
 Spatial Distributions of Tigers and Prey: Mapping and the Use of GIS
A map produced from a GIS is no better than the quality of the data that went into the GIS.
As with any computer-based system, garbage in produces garbage out! Hence, it is important
to incorporate high quality data with appropriate levels of accuracy for any GIS analysis.
Consideration should be given to various types of errors that may be present in the data
within a GIS. Positional accuracy can vary greatly among data sources, often simply due to
differences in the map scale originally used to create the data. Temporal accuracy often is a
problem; many types of GIS data quickly become out-of-date. The accuracy of content, or
thematic accuracy also may be a problem; the data may be coded, classified, or generalized
inappropriately for the goals of a particular project.
COMPILING SURVEY DATA IN THE GIS
GIS provides a centralized computer system for entering the data from the field survey forms
and associating those data with their proper geographic locations. The starting point for
entering data from the survey sheets is an accurate location of the area surveyed. Each survey
form must include location information; otherwise it is impossible to map the data. Ultimately,
all data within the GIS must be placed within some sort of map projection of cartesian
coordinates, or latitudes and longitudes. Therefore, there must be enough information on the
field survey forms and maps to enable the GIS analyst to properly georeference the survey
data within the GIS. The more information you provide by georeferencing - by marking on
your map accurately the latitude and longitude, or specific geographic features (villages,
rivers, etc.) that provide reference points - the more accurate your results will be.
The approach we will use to enter the survey data begins by generating polygons, which
represent the surveyed grid cells. Figure 6.1 shows a map of grid cells that might correspond
to areas surveyed for tigers and prey. To create the cells, the GIS technician enters a corner
latitude and longitude and dimension for the survey grids. Once a grid cell is created, clicking
on the grid cell with the computer mouse brings up a table that displays the data fields to be
filled in from the survey data form. The GIS collates the data from each grid cell into a large
table that can be queried or analyzed statistically (see below).
ANCILLARY SOURCES OF GIS DATA
In addition to the field data supplied from the survey forms, other sources of data will be
incorporated into the GIS to assist in map making and data analysis. These include road
networks, water bodies, political boundaries, locations of villages and cities, etc. Some of
these data layers are obtainable from state, national, or international agencies (e.g., Remote
52
Spatial Distributions of Tigers and Prey: Mapping and the Use of GIS
Sensing Agencies, Survey departments, Digital Chart of the World, etc.) in digital form that
can be integrated easily with the field data. Most digital data will already have an associated
map projection, but it is important to ensure that any data obtained have a map projection
that is compatible with the GIS software.
In some cases, the digital data may not be georeferenced, and ground control points will have
to be obtained to accurately georeference the data. It is also important to determine roughly
the spatial accuracy of the data you are obtaining. Data sets that are created from crude,
large-scale maps (e.g., 1:1,000,000) may be too inaccurate to be useful. For surveys of tiger
distribution, we recommend use of data from maps of scale 1:250,000 or finer resolution.
Satellite imagery and scanned aerial photographs can also be obtained from national or
international sources (e.g., IRS, SPOT, LANDSAT). When displayed on the computer screen,
these images provide a background to digitize important features directly on the computer
screen using a mouse, a process known as “heads-up” digitizing. Satellite imagery is useful
for coarse mapping of forest cover, and may be especially useful for updating older maps.
Aerial photography, if available, can be used to map forest cover with greater resolution and
accuracy.
In some instances, digital data will not be available and data must be digitized from hard
copies of maps or prints from satellite imagery using a digitizing tablet. The process of
digitizing by hand is very laborious and error-prone, but often it will be the only way to
incorporate certain types of data into a GIS. To successfully digitize from hard copy, at least
four, and preferably more, ground control points are needed to properly georeference the
digitized information.
SUMMARIZING DATA AND QUERYING THE GIS DATABASE
Although GIS is often viewed simply as a way to make maps, much useful information can be
produced from a GIS without creating a map. Summary statistics, tables of numbers, and
graphs can all be produced from GIS. In fact, many of the summary and query capabilities of
GIS are very similar to standard spreadsheet or database programs, since the underlying
database engines may be the same. Thus, apart from its map production capabilities, GIS is
a powerful tool for summarizing data and answering a wide variety of questions.
For example, using the summary statistic options of GIS, we can quickly determine what
proportion of the area with potential tiger habitat was not covered during the surveys. We can
count up the number of grid cells where tiger presence is detected and determine the proportion
53
 Spatial Distributions of Tigers and Prey: Mapping and the Use of GIS
of cells known to be occupied by tigers. We can then refine this information by asking how
many of the cells have known breeding populations of tigers, since these will be the cells that
are population sources. Other cells with tiger presence, but with no cub production, are likely
to be population sinks. Data can be cross-tabulated by state or region, allowing comparison
of the source-sink structure of different tiger populations. These examples illustrate only a
fraction of the types of questions that can be answered from summary statistics of the survey
data.
The tabulating capabilities of GIS also can be used for quality control of the GIS data. For
example, before producing a map, it is often a good idea to generate summary statistics for
variables that are going to be plotted. Frequently, such summaries will reveal missing or
erroneous data that should be fixed before making the map.
By comparing survey results from multiple years, we can look at crude trends in range
expansion and range contraction, for the establishment of new populations, for potential
dispersal routes, etc.
PRODUCING MAPS OF SPATIAL DISTRIBUTION
A map, like a picture, is worth a thousand words. Maps are invaluable tools for revealing
visual patterns that often are not evident from simple tables of numbers. In the past, map
production has been done laboriously by hand, requiring many hours to produce a single
map. GIS allows maps to be produced much more efficiently and inexpensively. With GIS it
is a simple matter to produce maps at different scales, covering different areas, and with
different color combinations or symbol sets. These advantages of GIS as a drafting tool are
considerable, but GIS can go beyond conventional maps to produce “analytical” maps that
are too difficult or impossible to create by hand. GIS can also be used to portray information
that could be generated by spatial analyses, which would otherwise not be evident from
conventional maps.
In Figure 6.2, we provide a simple illustration. From the survey data, we want to create a
map showing where breeding populations of tigers occur, and distinguish those areas from
areas with tigers that have little or no reproduction. This (hypothetical) source-sink map
reveals areas in need of some sort of management intervention, in the form of anti-poaching
efforts or habitat consolidation, etc. These sink areas may reveal important dispersal routes
used by tigers dispersing out of the source areas.
In subsequent chapters, we discuss methods of measuring relative or absolute abundance of
54
Spatial Distributions of Tigers and Prey: Mapping and the Use of GIS
tigers and prey species. Such data can then be used by the GIS to interpolate across unsurveyed
areas to create continuous surfaces that model abundance, using a process known as Kriging.
Figure 6.3 shows a map of hypothetical chital abundance in a park developed from point
estimates of density.
We can use maps such as Figures 6.2 and 6.3 to look for patterns of human use/impact and
their relationship with tiger presence, reproductive success, and prey abundance. Such analyses
will allow us to evaluate and improve our management strategies for patrolling, fire control,
and regulating other forms of human use. They also may provide data needed to spatially
model the population dynamics of tigers and prey.
ANALYZING TIGER-PREY-ENVIRONMENT RELATIONSHIPS
GIS provides a valuable mechanism for building a database that can be used to statistically
analyze the relationships between tigers, prey, and other environmental variables. This is the
final step in extracting information from the survey data, using multivariate statistical
techniques to look for relationships among these spatial variables. Although maps can show
many interesting patterns, maps can only display a few variables or relationships at a time.
To look at really complex relationships involving many variables, we will rely on multiple
regression techniques. These statistical techniques have the additional benefit of quantifying
the relationships among variables, allowing us to test hypotheses and make quantitative
predictions.
One useful approach, known as logistic regression, can evaluate which variables are the best
predictor of tiger presence-absence. If successful, this technique produces a simple equation
that shows the relative importance of different variables to tiger presence-absence. This equation
also shows us whether some of our hypotheses, for example, whether prey density is a good
predictor of tiger presence, are supported by the field data. If sufficiently robust, these statistical
models can also be used to predict the outcome of different management scenarios.
REFERENCES
Burrough, P.A. and McDonnell, R.A. 1998. Principles of Geographical Information System.
Oxford University Press, Oxford, UK.
Corsi, F., De Leeuw, J. and Skidmore, A. 2000. Modeling species distribution with GIS.
Pages 289-434 in Research Techniques in Animal Ecology (Eds: L. Boitani and T.
Fuller). Columbia University Press, New York, NY, USA.
55
 Spatial Distributions of Tigers and Prey: Mapping and the Use of GIS Spatial Distributions of Tigers and Prey: Mapping and the Use of GIS

Koeln, G., Cowardin, L. and Strong, L. 1994. Geographic Information Systems. Pages 540- Figure 6.1. Survey grid cells overlaid on potential tiger habitat. Cells are coded to show
566 in Research and Management Techniques for Wildlife and Habitats. Fifth edition presence/absence of tigers, and evidence of breeding.
(Ed: T. Bookhout). The Wildlife Society, Bethesda, MD, USA.

Morrison, M., Marcot, B. and Mannan, R. 1998. Wildlife-Habitat Relationships. Second

edition. University of Wisconsin Press, Madison, WI, USA.

Star, J. and Estes, J. 1990. Geographic information systems: An introduction. Princeton-

Hall, Inc., Englewood Cliffs, NJ, USA.

56 57
 Spatial Distributions of Tigers and Prey: Mapping and the Use of GIS Spatial Distributions of Tigers and Prey: Mapping and the Use of GIS

Figure 6.2. Map showing source-sink structure of tiger populations. Figure 6.3. Continuous surface model of chital abundance developed from point data samples.

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58 59
 CHAPTER 7

STATISTICAL CONCEPTS: INDICES OF RELATIVE

ABUNDANCE

James D. Nichols1 and K. Ullas Karanth2

United States Geological Survey, Patuxent Wildlife Research Center, Laurel, MD-20708-4017, USA.
1

Email: <jim_nichols@usgs.gov>

Wildlife Conservation Society (International Programs), Bronx, New York, NY-10460-1099, USA.
2

Email: <ukaranth@wcs.org>

KINDS OF INDICES

An index can be defined simply as “a measurable correlative of abundance” (Caughley 1977).

Based on the conceptual framework presented in Chapter 3, we can define an index as a count
BLANK statistic that is thought to convey information about abundance. Examples of possible indices
to abundance of tigers may include the number of tiger scats, tracks, or camera trapped
animals, or the number of kilometers of trail walked between successive encounters of tiger
sign. For prey species, possible indices could include the number of animals seen while walking
along a transect line, driving a stretch of road, or the number of piles of dung counted on a
plot.

Indices are typically used to draw inferences about relative abundances of animals over time
or space. Collection of the extra information needed to estimate p (equations 3.2 and 3.4) for
a particular count statistic can require substantial effort and cost, whereas use of simple
count statistics as indices is typically much easier and less expensive. Thus, use of indices to
estimate relative abundance is an attractive proposition to managers.

The utility of indices (e.g., specified as the strength of inference about relative abundance)
depends strongly on the nature of the relationship between the index and true abundance and
on our knowledge of this relationship. A monotonic relationship between index and abundance
is essentially a relationship that is always in the same direction; thus if there is a positive
relationship between the index and average abundance then, on average, a larger index value
means a greater abundance. A monotonic relationship is usually assumed in studies or
monitoring programs that use indices and permit general inequality statements about abundance
at different times or locations. For example, let CA and CB be count statistics for locations A
and B that are monotonically (and positively) related to animal abundance. Then if average
counts at A are greater than those at B [i.e., E(CA ) > E(CB )], we might conclude that abundance
is also greater at A than B (NA > NB ).

61
 Statistical Concepts: Indices of Relative Abundance
However, there are two problems with drawing inferences about abundance (N ) from index
statistics, even when we know the relationship between index and abundance to be monotonic.
First, count statistics are random variables, and we do not know E(CA) or E(CB). Instead, as
suggested above, we must use some sort of replication-based approach to compute average
counts (averages frequently provide good estimates for expected values) for the two places or
times being compared. We must also obtain information (e.g., via the replication on which the
averages are based) on the variances of the two count statistics in order to even make an
inequality inference.
For example, assume that we are interested in chital abundance in two different areas (denoted
as A and B) of habitat and have a prediction about abundance being greater in one habitat
type than the other. We might choose to index chital abundance by dung counts in randomly
selected sample plots from the two areas. Say that we count dung piles in 10 different plots
(size 100 m2 each) in each habitat, and compute the average number of piles per plot (denoted
in area A as C A ), and associated standard error (denoted as SEˆ (C A ) ), for each habitat (each
set of 10 plots). If CAi denotes the count in sample plot i of area A, then the average (or mean)
and standard error are computed as:
n
∑C Ai
CA = i =1
n or places really do indicate different numbers of animals. This problem is more difficult, as it
n investigators (it is not dealt with).
∑ (C Ai − CA )2
SEˆ (C A ) = i =1
n( n − 1)
where n=10 is the number of sample plots at which dung piles are counted. Finally, note that
an approximate 95% confidence interval for C A can be constructed as:
C A − 1.96 SEˆ (C A ), C A + 1.96 SEˆ (C A ).
Assume that we construct approximate 95% confidence intervals from the standard errors
and obtain the values of:
C A = 2.1, 95% CI = 1.7, 2.5 and C B = 1.6, 95% CI = 1.3, 1.9
We could construct a t test directly from the plot counts (e.g., Steel and Torrie 1960), or we
can examine the degree of overlap of the confidence intervals of the average counts. Using the
latter approach, we see that the overlapping confidence intervals for the two areas, A and B,
62
Statistical Concepts: Indices of Relative Abundance
indicates that we do not have strong evidence that the number of dung piles in the two areas
is really different.
The second problem with drawing inferences about abundance from index statistics,
even when we know the relationship between index and abundance to be monotonic, involves
the strength of the relationship between abundance and the count statistic and the uncertainty
associated with our estimation of the relationship. This relationship will not be perfect in
the sense that there will be error associated with any prediction. Assume that an average
count of two piles per 5 m2 corresponds to a prediction of 40 chital per km2. If we
obtain an average of two piles per 5 m2 based on random samples within an area of
interest, we would not expect exactly 40 chital per km2 in this area, but instead would realize
that there would be error associated with our prediction such that we might actually have 36
chital per km2, or perhaps 43. Therefore without actually estimating the relationship (e.g.,
from count data and actual abundance estimates), the magnitude of this variation will be
unknown.
Thus, the two problems are associated with uncertainty. The first problem involves variation
in the count statistics themselves, and this problem can be dealt with via replication, permitting
inference about whether the index really does differ between the times or places being compared.
The second problem involves uncertainty about whether different index statistics in two times
involves some sort of calibration of the index, and is usually assumed away by many
A linear relationship is a special kind of monotonic relationship of the form:
(C = β 0 + β1 N + ε ), where β0 is an intercept parameter, β1 is a slope parameter, and ε is an
error term characterizing the uncertainty and variation of the relationship. A linear relationship
permits inequality statements, although inference is again dependent on the ability to estimate
variances of counts (e. g., via replication) and on some knowledge of the error term of the
relationship.
A special case of the linear relationship is a direct proportional relationship in which
the index is directly proportional to true abundance: (C = pN + ε ) , based on the notation
of equation 3.1. An index showing such a relationship with abundance is termed a proportional
index or constant-proportion index (Lancia et al. 1994) and permits direct inference
about relative abundance. This relationship is commonly assumed in the use of index
statistics.
Here we consider the use of proportional indices to draw inferences about relative abundance.
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 Statistical Concepts: Indices of Relative Abundance
Define relative abundance for two places or times, 1 and 2 as:
λ = N 2 / N1,
where N i denotes abundance at time or place i. If abundance is directly estimated for each
time or place, then we can estimate relative abundance as:
λˆ = Nˆ 2 / Nˆ 1 (7.1)
If we have only count statistics, we may still be able to estimate relative abundance if these
index statistics meet the conditions of a proportional index. Consider the estimator.
λˆ = C 2 / C1
The expected value of this estimator can be approximated as:
E (C 2 ) p N
E (λˆ ) ≈ = 2 2
E (C1 ) p1 N1
where p1 and p2 are the two proportionality constants (from equation 3.1) relating the index
N survey or monitoring programs (see Conroy and Nichols 1996). The ability to apply these
statistic to true abundance. Note that E (λˆ ) ≈ 2 = λ ( i.e., λˆ is approximately unbiased),
N1
only when p1 = p2. This is a very restrictive condition and thus represents a strong assumption.
If the detection probabilities themselves are considered to be random variables from some
distribution, then the ratio of index statistics provides a reasonable estimate of λ only when
E(p1) = E(p2). Generally, this latter condition requires that the distribution of detection
probabilities remains the same over the times or places being compared. Again, this represents
a strong assumption that should be investigated if index statistics are to be used to estimate
relative abundance.
As an example, assume that two areas have exactly the same number of chital deer: N1 = N2.
Further assume that we use the number of deer seen while walking one km of transect as an
index to deer abundance, and that 100 animals are potentially observable from the transects
in each habitat. Habitat differences between areas 1 and 2 cause the detection probability for
deer in area 1 to be p1 = 0.20 and the detection probability for deer in area 2 (a more dense,
heavily wooded area) to be only p2 = 0.15. Then, on average, we will see about 20 animals per
km transect walked in area 1 (E(C1) = 20) and about 15 deer per km transect in area 2 (E(C2)
= 15). So the expected value of our estimate of relative abundance based on count statistics
treated as indices is:
64
Statistical Concepts: Indices of Relative Abundance
E (C 2 ) 15
E (λˆ ) ≈ = = 0.75
E (C1 ) 20
Our index-based conclusion would be that area 2 contained only about 75% of the chital
N
present in area 1. True relative abundance in this case is: λ = 2 = 1.0 . Bias of this
N1
relative abundance estimator is then given by the difference between the expected value of the
estimator and the true value of the quantity being estimated:
E ( λˆ ) − λ = 0 . 75 − 1 . 00 = − 0 . 25 = bias of λˆ
REDUCING BIAS IN INDEX-BASED ESTIMATES
Use of count statistics as indices to draw inference about relative abundance is thus based on
the assumption of equal detection probabilities (or, more generally, equal proportionality
constants relating the index to true abundance) for the times or locations being compared.
Investigators who use indices for estimation of relative abundance must thus try to reduce
differences in p. There are different approaches to dealing with detectability in index-based
different approaches depends on our ability to identify and influence the sources of variation
in detection probabilities and index proportionality constants.
The first approach is standardization and simply involves use of standardized methods to try
to produce similar detection probabilities or proportionality constants for the areas or times
being compared. This approach can be effective for factors influencing p which the investigator
can identify, and over which the investigator has some control.
A frequent example of standardization involves attempts to expend the same amount of effort
at the different times and places to be compared. Effort can be quantified in many different
ways and might involve the amount of time spent searching or counting, the linear distance
traveled during search or count efforts, the amount of area searched, the number of person-
hours expended, the number of traps set, etc. Use of persons with similar training and abilities
to detect animals also represents an attempt to equalize effort. In addition, clear written
instructions and training can be important components of standardization when different
people (as opposed to the same person) are involved in the monitoring effort. Another kind of
standardization involves conducting surveys or counts at similar times of the day, during the
same season of the year, and under similar weather conditions (e.g., it is common to set limits
of temperature or rainfall beyond which counts are not to be conducted).
65
 Statistical Concepts: Indices of Relative Abundance
Another approach to reducing bias associated with variation in detectability involves use of
covariates believed to be associated with this variation. Sometimes investigators can identify
factors that influence p, but that cannot be readily controlled. In such cases, covariate
information can be collected and used in the analysis and interpretation of index statistics. An
important requirement for the reasonable use of covariates with indices is that they are
associated with variation in p but not with variation in true abundance.
Examples of covariates include environmental covariates at the time of the survey
(e.g., ambient temperature during line transect surveys of deer or substrate condition during
dung/track surveys). Note that some of these factors can be dealt with via standardization
as well. The appropriate data-analytic method for dealing with such covariates is often
analysis of covariance (ANCOVA; see Steel and Torrie 1960). In some cases, work is done
to model the relationship between p and the covariate explicitly, and in such cases
estimates of p are essentially obtained and incorporated directly into the analysis (regarding
terminology, we are essentially using an estimation, rather than an index, approach in such
situations).
Effort is sometimes used as a covariate (e.g., numbers of trap nights, linear distance traveled,
area searched, hours searched, etc.). Again, such covariates can also be dealt with via
standardization. Data-analytic methods for dealing with effort covariates include ANCOVA
and creation of new effort-adjusted count statistics (e.g., animals caught per trap night, animals
seen per km traveled, animals seen per search hour). In cases where p can be modeled explicitly
as a function of effort we essentially move from use of indices to use of estimation methods
such as removal and catch-effort models (Seber 1982; Williams et al. 2002).
Habitat covariates can potentially be incorporated into analyses of index data (e.g., visibility
will be a function of vegetation density as in dense understory vs. short grass savannah).
Sometimes ANCOVA is used to deal with habitat covariates. This approach, however, has
the potential to be misleading. Recall that E(C) = Np (equation 3.1), and habitat likely influences
both N and p in many situations. It is thus important when dealing with covariates in index
surveys to be sure that the covariate is relevant only to detection probability (or the
proportionality constant) and not to abundance also.
Unfortunately, there are likely to be many factors that influence detection probability (and
proportionality constants) that we cannot identify or control. When using indices, we must
simply hope that such variation is relatively small and not misleading. Surveys based on
indices thus yield weak inferences in the sense that results and conclusions are always tentative;
that is they are always based on untested assumptions. Thus, if we observe that an index
shows decline over time, we can never be certain whether the decline is in the quantity of
66
Statistical Concepts: Indices of Relative Abundance
interest, animal abundance, or in the nuisance parameter, p, the proportionality term relating
the index statistic to abundance.
ROBUST ESTIMATION OF RELATIVE ABUNDANCE
The above approaches to minimizing bias in estimates of relative abundance (standardization,
covariates) are important for use of indices, but are not nearly so important for use of formal
estimation methods (e.g., those described in Seber 1982; Williams et al. 2002). When p is
directly estimated, then unbiased estimates of relative abundance can be obtained in the absence
of standardization and covariate measurement, as these are dealt with via estimation of
potentially different p’s for each sampling location or occasion. Similarly, there is no need to
identify all the factors that influence detection probability. However, standardization still
offers advantages even in these situations, as it becomes possible to test for differences in p
and, if no differences are found, to use the count statistics to develop estimates of relative
abundance that have smaller variances than estimates based on Nˆ i .
Skalski and Robson (1992) recommend the following approach to estimating relative abundance
when the information necessary to estimate p is collected. They suggest first testing for equality
of detection probabilities for the two locations or occasions being compared (i.e., test H0, that
p1 = p2). If there is evidence that the detection probabilities differ (i.e., if H0, is rejected), then
relative abundance should be estimated as the ratio of abundance estimates:
Nˆ
λˆ = 2
Nˆ 1
(as in equation 7.1).
However, if the test of p1 = p2 provides no evidence of differences in detection probability,
then relative abundance can be estimated as:
C
λˆ = 2 (7.2)
C1
Equation 7.2 simply estimates relative abundance as the ratio of count statistics. The estimator
of equation 7.2 will tend to have a smaller variance than that of equation 7.1, because equation
7.1 requires the estimation of p1 and p2 in order to estimate abundance. This extra
variance component associated with the estimation of detection probability is not
included in the estimator of equation 7.2.
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 Statistical Concepts: Indices of Relative Abundance
SUMMARY COMMENTS ON INDICES AND ESTIMATION
Indices typically require much less cost and effort than direct abundance estimation methods
and can be used to draw inferences about relative abundance. However, the “cost” associated
with indices is that their use is grounded in faith that untested assumptions are not violated; in
particular that p1 = p2, or at least that E(p1) = E(p2). Indices will be most useful when this
faith is justified or can be defended. At a minimum, use of indices to estimate relative abundance
requires vigilance for differences in p’s and an appreciation for the bias that can be produced
by such differences.
We believe that use of indices to estimate relative abundance over time (“trend”) is likely to
be safer (result in less biased estimates) than use of indices to estimate relative abundance
over space. Habitat and site-specific differences in detection probability are typically less
important when focusing on temporal change in the same locations than when investigating
changes in abundance from one location to another.
The best approach to estimating relative abundance is to collect the information needed to
estimate detection probability simultaneously with the collection of the count statistics
themselves. This approach carries the additional cost and effort of this extra data collection.
However, the benefit of this approach is that it is possible to: (1) formally test hypotheses
about difference in detection probabilities between the areas or times under comparison and,
if evidence of differences is found, (2) estimate relative abundance in a manner that properly
incorporates these differences.
The decision about whether to use indices or estimation methods in animal monitoring
programs is ultimately a decision about the relative importance of two costs: The cost
of the monitoring program itself versus the cost of uncertainty and weak inference. Our
basic recommendation is that if a biological or management question is sufficiently
important to merit the effort associated with design and conduct of an animal population
monitoring program, then it is probably also sufficiently important that estimation be taken
seriously. We recognize that collection of the additional information needed for proper
estimation is simply not possible in some situations, but we believe that these situations
are much rarer than commonly portrayed. More detailed discussions of monitoring
methods that reflect the perspective presented here can be found in Thompson et al. (1998),
Nichols et al. (2000), Yoccoz et al. (2001), and Pollock et al. (2002).
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Statistical Concepts: Indices of Relative Abundance
REFERENCES
Caughley, G. 1977. Analysis of Vertebrate Populations. Wiley, New York, NY, USA.
Conroy, M.J. and Nichols, J.D. 1996. Designing a study to assess mammalian diversity.
Pages 41-49 in Measuring and monitoring biological diversity: Standard methods
for mammals (Eds: D.E. Wilson, F.R. Cole, J.D. Nichols, R. Rudran and M.S. Foster).
Smithsonian Institution Press, Washington, DC, USA.
Lancia, R.A., Nichols, J.D. and Pollock, K.N. 1994. Estimation of number of animals in
wildlife populations. Pages 215-253 in Research and management techniques for
wildlife and habitats (Ed: T. Bookhout ). The Wildlife Society, Bethesda, MD, USA.
Nichols, J.D., Hines, J.E., Sauer, J.R., Fallon, F.W., Fallon, J.E. and Heglund, P.J. 2000. A
double-observer approach for estimating detection probability and abundance from
point counts. Auk 117:393-408.
Pollock, K.H., Nichols, J.D., Simons, T.R. and Sauer, J.R. 2002. Large scale wildlife
monitoring studies: Statistical methods for design and analysis. Environmetrics (in
press).
Seber, G.A.F. 1982. The estimation of animal abundance and related parameters. Second
edition. MacMillan, New York, NY, USA.
Skalski, J.R. and Robson, D.S. 1992. Techniques for Wildlife Investigations. Academic
Press, San Diego, CA, USA.
Steel, R.G.D. and Torrie, J.H. 1960. Principles and procedures of statistics. McGraw-Hill,
New York, NY, USA.
Thompson, W.L., White, G.C. and Gowan, C. 1998. Monitoring vertebrate populations.
Academic Press, New York, NY, USA.
Williams, B.K., Nichols, J.D. and Conroy, M.J. 2002. Analysis and management of animal
populations. Academic Press, San Diego, CA, USA.
Yoccoz, N.G., Nichols, J.D. and Boulinier,T. 2001. Monitoring of biological diversity in
space and time. Trends in Ecology and Evolution 16:446-453.
69
 CHAPTER 8

FIELD SURVEYS: ASSESSING RELATIVE ABUNDANCES OF

TIGERS AND PREY

K. Ullas Karanth1 and N. Samba Kumar2

1
Wildlife Conservation Society (International Programs), Bronx, New York, NY-10460-1099, USA.
Email: <ukaranth@wcs.org>
2
Centre for Wildlife Studies, 26-2 Aga Abbas Ali road (Apt. 403), Bangalore, Karnataka-560 042, India.
Email: <wcsind@bgl.vsnl.net.in>

INDEX SURVEYS

Ideally, estimates of absolute abundances of tiger and prey species derived using the approaches
described in Chapters 9-12 provide the best means for reliably monitoring populations of
BLANK these animals. However, full-blown line transect or capture-recapture surveys require skilled
manpower, appropriate equipment and substantial financial resources. These skills and
resources may not be available in many field situations. In such cases, for monitoring tiger
and prey populations in individual wildlife reserves or study sites, we can try to use indices of
relative abundance conceptualized in the framework presented in Chapters 3 and 7.

The objective of these types of sample surveys is to generate quantitative indices of abundance
that are positively correlated with the true densities of tigers and prey species in the area.
Such quantitative indices will possess estimates of mean values as well as the standard errors,
confidence intervals and coefficients of variation associated with them (Chapter 7). If the
indices of tiger or prey abundance can at least tell you whether a given tiger or prey population
has increased, decreased or remained stable between surveys, they would be quite useful for
assessing the value of management interventions. When deriving such indices of relative
density we hope that the relationship between the true abundance of animals and the index
used is strong (Chapter 7).

We can of course never be certain that the indices are predictably related to the actual abundance
of tigers and prey (Shenk et al. 1998; Yoccoz et al. 2001). We hope that by standardizing the
survey methods in terms of variables that affect our counts, such as weather, survey effort,
area surveyed, season, etc., we can derive ‘constant proportion indices’ described in Chapter
7 where the value of p remains unchanged between two sample surveys. Therefore, the proper
design of the sample survey and the choice of the right type of field measurement are both
crucial in the task of deriving a useful index of abundance for tigers or prey.

71
 Field Surveys: Assessing Relative Abundances of Tigers and Prey Field Surveys: Assessing Relative Abundances of Tigers and Prey

An adequate index survey will involve substantial field effort to collect large samples of data
to generate a more precise index. The survey should also representatively cover the entire
area of interest for the index to truly reflect the status of the tiger or prey populations within
it. In the following sections, we discuss some possible field methods for index-based surveys
of tiger and prey species, with brief comments on their potential usefulness and limitations.
INDICES OF RELATIVE ABUNDANCE FOR TIGERS
Indices from Encounter Rate Surveys of Tiger Sign
Tigers prefer to use roads or trails made by other animals as travel routes (Smith et al. 1989;
Karanth and Nichols 2000). In most places, tiger signs like tracks and scats are more likely to
be found on such travel routes. However, substrate conditions vary widely across the vast
distributional range of tigers. Clear tiger tracks are particularly difficult to find unless the
substrate comprises of dust, sand, mud or snow. Such suitable substrates occur in very few
sites where the natural soil type – in combination with vehicular traffic – creates ‘track plots’
along roads on which tiger footprints are imprinted clearly.
recommend choosing 40-50 such ‘sampling routes’. However, if such predetermined sampling
routes cannot be used, survey teams can also traverse trails/routes on the day of the survey
itself, using pedometers, maps, and compasses or a global positioning system (GPS), to keep
track of the survey routes used and the sampling effort invested.
In order to obtain a precise index of tiger abundance, large samples of data are required. The
total sampling efforts invested in surveys of tiger scat and tracks will depend on the manpower
resources available at the site. For example, if you have identified 40 road segments of 15 km
length each, they add up to a total sampling effort of 600 km. If four teams, each covering 15
km/day are deployed, the survey can be completed in 10 days; if eight teams are deployed the
same distance can be covered in five days.
If the same sampling routes are replicated several times over a period in order to estimate
detection probabilities for signs, the number of days in the interval between two successive
sampling efforts has to be long enough for the tracks or scats encountered in the earlier
survey to disappear completely. Another approach is to remove the detected sign (collect the
scat, brush away the tracks) during the conduct of each survey.

Before track surveys are carried out, definition of an encounter (a data point in the survey)
Artificially creating suitable track plots has been suggested as a solution to the widespread
has to be established clearly. For example, if the observers follow the same set of tiger tracks
problem of absence of appropriate substrates. However, we have observed that such artificial
for several hundred meters or even several kilometers, should it be defined as a single encounter?
track plots are difficult to create or maintain, and they rarely work efficiently in practice.
Some predetermined criteria have to be established to deal with such problems. One criterion
Lack of suitable soil in the vicinity, logistical problems in transporting soil, and the effect of
may be that only the first encounter with a continuous set of tiger tracks is recorded. Alternately,
rain, wind, and animal movements, often hinder the use of track plots under field conditions.
encounter with the same set of tracks after a ‘break’ (caused by poor tracking substrate or the
animal moving off the trail and then returning to it), can be recorded as a new encounter.
Tigers generate more tracks than scats per day, and generally more tracks are seen than scats.
After a consistent definition of a ‘track encounter’ is established beforehand, the index estimate
However, if the substrate consists of grass, leaf litter or hard soil, scats will be easier to detect
of tiger density will be the number of ‘encounters’ per 10 kilometers (or some other unit of
than tracks. Therefore, we suggest carrying out simultaneous surveys of tiger tracks and
distance) walked by the survey teams (e.g., 5.6 tiger tracks/10 km).
scats, and, finally using whichever sign that yields a better index. In presence-absence surveys
designed to estimate the proportion of area occupied, both scats and tracks are indicators of
An approach that avoids the problem of defining a ‘track/scat encounter’ is to mark segments
tiger presence. In cases where both tracks and scats are encountered regularly during sampling,
of trails or roads to be surveyed at 0.5 or 1.0 km intervals. Then, if survey teams walk a total
it may be useful to consider a combined index in situations where relative abundance is the
of 600 kilometers, they cover 1200 sampling units of 0.5 km length or 600 sampling units of
parameter of interest.
1.0 km length. In this case, the observer simply records whether tiger tracks were present in
a sampling unit. The index of tiger abundance in this case is the proportion of the sampling
Before designing the survey, a suitable network of roads and trails should be identified in the
segments in which tiger tracks are encountered. For example, an index estimate of relative
study area. This network should cover the area to be monitored representatively (Thompson
tiger abundance would be 0.22, indicating that 22% of all sampled segments had tiger tracks
1992; Thompson et al. 1998). A sampling scheme should then be designed (Chapter 5) in
present in them.
which several different road/trail segments, each one of about 10-15 km length (a distance
that can be walked slowly by a survey team during one day’s work) should be identified. We
Defining an encounter is easier with tiger scats than with tracks, because the former are

72 73
 Field Surveys: Assessing Relative Abundances of Tigers and Prey Field Surveys: Assessing Relative Abundances of Tigers and Prey

discrete entities. In a scat index survey, for example, the estimated number of tiger scats recording the number of scats encountered during sample walks, it is useful to measure the
encountered/10 km walked, or the proportion of trail segments that contained a tiger scat diameter of the scat and collect the specimen in a plastic cover, after individually numbering
would be the index generated. Tables 8.1 and 8.2 illustrate results of index surveys of tiger it. The scats can later be used for studying tiger food habitats through dietary analysis.
scats conducted in five reserves in India during 1995-1996 (K.U. Karanth, unpublished data).
It may even be possible to extract the minute amount of the tiger’s DNA present in the scat
Field surveys should be conducted during the season when tiger tracks are easily found. The and use it to identify individual tigers. Such individual identities can then be used in combination
surveys should be started early in the morning before other wild animals, vehicles, people, with advanced capture-recapture population estimation protocols covered in Chapters 11 and
livestock and wind begin to obliterate tiger signs. 12. Care should be taken while collecting scats: It is crucial that specimens do not get
contaminated during collection and they should be uniquely labeled and stored. Protocols for
A team of at least two observers should walk each sampling route, carefully searching the collection of scats for DNA analysis are described elsewhere (Wasser et al. 1997; Ernest
road/trail for tiger tracks on either side (Figure 5.1). On encountering a tiger scat or a set of et al. 2000).
tracks the team should record the event as well as the time, location, measurements of size
etc. The team should also record the starting and ending time and locations for each sample Index Surveys of Tigers Using Camera Traps
walk. In areas where tiger tracks are scarce or likely to be confused with those of leopards, or
At many sites, it may be impossible to survey tiger sign due to constraints such as lack of
‘voucher’ specimens are needed for better supervision, it may be necessary to collect plaster
enough personnel, logistical problems, poor substrate or weather conditions. In such situations,
casts or photos of the tracks.
sometimes, self-activated camera traps can be deployed to photograph tigers. Details of
conducting camera trap surveys and their use for estimating absolute densities of tigers are
To keep track of the sampling effort, it is essential to accurately record distances covered
provided later in Chapters 11-13. Here we restrict ourselves to describing the potential use of
during each sample walk. The distance traveled can be measured using a pedometer or a GPS
camera traps for deriving indices of tiger abundance based on trapping rates (e.g., number of
unit during the survey. It can also be measured using a tape, from a map, or using the odometer
tiger photos/100 trap nights of effort).
on a vehicle, prior to the survey.
In the case of camera trap surveys, the sampling effort is measured in terms of the number of
In the mangrove forests of the riparian delta in Sundarbans of India and Bangladesh, carrying
camera trap nights. Each trap night represents a trapping effort over one day (either a 12-
out track surveys on land as described above poses unique problems. The presence of man-
hour period from dusk to dawn if cameras are put out only at night, or, the entire 24 hour
eating tigers precludes extensive sign surveys on foot. The soft soil results in imprints of
cycle if cameras are always deployed). For example, five cameras deployed over 10 days
some old tracks being retained for a long time, whereas daily tidal patterns erase some other
result in 50 trap nights (or trap days) of effort.
tracks quickly, thus making it difficult to define a ‘track encounter’. To overcome these
problems, K.U. Karanth and J.L.D. Smith (unpublished data) designed a track survey method
Each tiger photograph obtained is then treated as an encounter. These surveys will generate a
specific to the Sundarbans mangrove habitat. In this method, surveys of tiger sign are carried
simple quantitative index, consisting of the number of “camera trap encounters” (photographic
out from boats that traverse the numerous, well-distributed creeks and canals that exist in the
captures) of tigers per unit sampling effort. For example, the index estimate may be something
area. The boats proceed at a pace of about 4.0 km/hour, while two observers carefully examine
like 2.4 tiger photo captures/100 trap nights of effort. An example of this is given in Table 8.3
the inter-tidal zone on the banks on either side of the boat. They count every ‘fresh’ set of
obtained from data collected at seven sites in India (K.U. Karanth, unpublished data).
tiger tracks that cross the canal. Because the daily tidal fluctuations obliterate old tracks on
the embankments, discrete encounters with fresh tracks are easy to define. From these boat
Although some workers claim that such raw trapping rates are also good predictors of absolute
surveys, the tiger abundance index is estimated as the number of track sets/10 km traveled by
densities of tigers in an area (Carbone et al. 2001), there are major statistical and ecological
boat.
arguments against accepting this premise (Jenelle et al. 2002; Chapter 7). Therefore, even
when the survey objective is only to derive an index of tiger abundance, we strongly recommend
An example of a field data form that can be used for scat-based index surveys of tiger density
camera trapping using the formal capture-recapture protocols described in Chapters 11-13.
is provided in Appendix 2.1. When tiger scats are counted during surveys, in addition to

74 75
 Field Surveys: Assessing Relative Abundances of Tigers and Prey
Even if such camera trapping data are too sparse to generate good estimates of absolute
abundance of tigers, capture-recapture sampling can still convey some ideas about underlying
capture probabilities. With this approach you can at least try to judge to what extent our
indices based on trapping rates are unbiased and robust, as described in Chapter 7. In addition,
it may be possible to use information from camera trap surveys in other areas to estimate
detection probabilities. The main point to emphasize is that camera trapping is a survey
approach that is relatively expensive. In most cases where the expense and effort associated
with camera trapping are justified, it makes sense to try to attain the strongest inferences
possible, and, these will typically result from the methods described in Chapters 11-13.
INDICES OF RELATIVE ABUNDANCE FOR PREY SPECIES
Most prey species can potentially be monitored using field methods that employ direct sightings
of animals. The estimation of absolute abundances of prey using such distance-sampling
methods is treated in Chapters 9 and 10. In this chapter, we look at indices of relative densities
for prey species, based on sightings. We note however, that in areas where prey species are
difficult to see due to wariness or other factors, we may be forced to use indices of relative
abundance based on signs such as dung or tracks rather than direct sightings.
Index of Encounter Rates from Line Transects
Line transects are straight, narrow walking trails that are cut so that they traverse the area
being surveyed according to some pre-planned sampling design. The advantage of such
transects is that they sample the surveyed area representatively so that inferences about animal
abundance made from them can be extrapolated to the larger area (Chapter 9).
The design of appropriate sampling schemes for placement of transects and field procedures
for carrying out line transect surveys, are described in Chapters 9 and 10, in the context of
formal line transect sampling. The only difference between such full-fledged line transect
surveys and the encounter rate surveys described here is that in the latter surveys, observers
only count the animals they encounter without collecting associated distance data. These
count data are then used to estimate an index of relative abundance. For example, a foot
survey in which 16 sambar were counted while walking 10 kms would yield a mean index
value of 1.6 sambar/km walked.
In the absence of estimation of sighting probabilities based on distance data, such encounter
rate-based indices are not very useful to compare prey abundances between different areas, if
sighting conditions (for example density of cover) are very different. They may be more
76
Field Surveys: Assessing Relative Abundances of Tigers and Prey
useful to compare temporal changes in prey abundance at the same site, if the detection
conditions remain unchanged between surveys.
We emphasize that formal line transect surveys (Chapters 9 and 10) are to be preferred over
encounter rate surveys, wherever possible. However, in situations where distance-measuring
instruments and compasses are not available, or there is a lack of skilled survey personnel
who can use such instruments, index-based encounter rate surveys can be adopted.
We are aware that some investigators have derived indices of ungulate abundance (or even
absolute abundance estimates) from counts made from vehicles while driving along roads.
Some others have used counts from observation platforms near water holes or other areas,
and, from ‘drive counts’ (Rodgers 1991). However, roads and observational platforms do
not representatively sample the abundance of prey animals within a surveyed area. Applying
formal distance sampling models (line transects and point transects) to such roadside or
platform count data is not justified because of the serious violations of the underlying model
assumptions (Chapter 9). In particular, any sort of orientation of animals with respect to the
point or transect from which observations are made can result in true density gradients that
have nothing to do with detectability and thus invalidate use of distance sampling estimation
methods. We are not aware of any effort to statistically model the data resulting from drive
counts, in which animals are chased by multiple observers and counted. Thus no estimators
have been derived for this method.
Therefore, we generally do not recommend roadside, platform and drive counts (Rodgers
1991) for monitoring the abundance of prey species.
Index of Prey Density from Camera Trap Surveys
Camera trap surveys offer yet another option to estimate relative abundance of prey species
in areas where animals are difficult to see. Camera traps can be deployed along animal trails
and paths to generate indices of prey density in terms of number of photographic captures per
unit trapping effort. For example, an abundance index derived from camera trapping for
ungulates may give a figure of 8.6 Muntjac photos/100 trap nights. However, the strength of
the relationship of the index with true prey abundance will remain unknown (Chapter 7) until
the index can be validated using an independent estimate of absolute density derived from
formal line transect surveys (Chapter 9).
Indices Based on Surveys of Animal Dung
Because dung (or pellets) of ungulate prey are often easier to find, dung counts are more
widely used to estimate their relative abundance. However, to derive useful indices of abundance
77
 Field Surveys: Assessing Relative Abundances of Tigers and Prey
from dung counts, deployment of a proper sampling scheme and the standardization of survey
protocols are essential.
The key consideration in dung surveys is that the plots representatively sample the surveyed
area (See also Chapters 3 and 4). The survey design should ensure that adequate sample sizes
are likely to be attained while counting dung piles in order to generate indices with sufficient
precision.
The entire area to be surveyed can be overlaid with a grid and the survey plots set out according
to some sampling design (Chapter 4). Alternatively, a line transect type survey design (Chapter
9) can be used. In the latter case, the dung plots are placed on the transect lines, at intervals
of 100 meters or so. Instead of laying the plots linearly on the transect line itself, along which
animals may sometimes walk, dung plots can be placed perpendicular to the transect orientation.
Although dung survey plots can be of any shape (circular, square etc.), we found rectangular
strips 50 × 2 meter size practical to use (Figure 8.1).
Figure 8.1 - Survey team counting animal dung on a 50 x 2 meter plot.
78
Field Surveys: Assessing Relative Abundances of Tigers and Prey
Survey personnel should be able to recognize and identify the dung or pellet of different
species. Prior training of survey personnel in dung identification is, therefore, critical. It is
also important to define what is to be counted as a ‘dung pile’ or a ‘pellet group’ clearly
before the survey. The key is to adopt a uniform survey protocol for the all personnel involved.
A survey team of three persons (two to count dung piles and the third to maintain the record)
should thoroughly search each plot and record all dung piles of prey species present inside
the sample plots. Finally a total count of all dung piles encountered is recorded for each
species.
For dung plots of 100 m2 size, surveyed by three persons, we suggest the following field
procedure, as an example: First, the dung plot is laid with the help of a 50-meter nylon rope
stretched out at each sample point. One observer searches on either side of the rope carefully,
holding a one-meter long stick perpendicular to the rope, as a marker. While counting dung
piles, the observers should take care not to trample on them. A third observer records their
counts within the plot. Each dung pile counted might consist of several individual boluses or
pellets. The survey goal usually is to count the number of piles, not of individual pellets or
boluses. After the dung counts are tabulated, the relative abundance index for each species of
prey is provided by the density of dung (e.g., 4.4 chital pellet groups/hectare; 2.8 gaur dung
piles/hectare). Appendix 2.2 shows a typical data form used for recording dung count data in
the field.
Estimating Dung Decay Rates
However, there is a major problem in using dung density as a valid index of prey abundance
in any area. When dung piles are counted for the first time, a count of ‘standing crop’ of dung
piles in the area is obtained. However, these dung piles would have been deposited over
several days prior to the count. Therefore, the counts record only the ‘survivors’, among the
piles that suffered a process of natural decay during days that preceded the count.
Therefore, if ‘standing crop’ dung counts are to be used to compare densities at two different
points in time (or between two different sites), we must be sure that both of the counts represent
dung deposited over comparable periods: They must both be counts of dung deposited, for
example, over the preceding one month.
One approach to meeting this assumption is to clear the plots completely of all old dung first
and then counting piles after the same interval at both sites. However, clearing plots before
doing the counts requires more manpower, effort and time than simply counting the standing
crop of dung. If these are constraints, prior clearing of plots is not a practical option.
79
 Field Surveys: Assessing Relative Abundances of Tigers and Prey
If comparisons are made from standing crop dung surveys, the daily rate of deposition of
dung piles (the number of dung piles arriving on the plot/day) must be estimated. To estimate
daily dung deposition rates, the number of piles that decayed (did not survive) before the
count must be estimated. Decay rates can potentially differ greatly between two sites or at
two different times. Therefore, estimating the decay rates that prevailed in the days prior to
the dung survey is critical (Hiby and Lovell 1991).
To measure dung decay rates before the counts, it is necessary to set up separate dung decay
experiments in advance of the actual survey (Hiby and Lovell 1991; Marques et al. 2001).
Fresh dung piles (<24 hours old) should be located and tagged with the dates on which they
were produced. Let us assume that, after an experiment as above, at one site we found that
chital dung piles decayed completely in 20 days, and at another site they took 30 days. Thus
the decay rates work out to 1/20 in the first site and 1/30 at the second site. Let us say we
counted dung to obtain a dung density of 400 piles/hectare in the first site and 600 piles/
hectare at the second site. Correcting for differences in decay rates, the dung arrival rate at
both sites work out to 20 piles/hectare/day. In other words, although we counted more pellets
in the second site, this was because of a slower decay rate, rather than due to any real difference
in chital density. However, in practice, dung decay rate experiments are not easy to perform
if dung density (animal density) is low.
Sophisticated approaches to dung decay rate estimation that include complex, but biologically
more realistic computations are available (Hiby and Lovell 1991; Marques et al. 2001). If
dung densities are to be used as indices of relative abundance of prey species, we strongly
recommend that these approaches be used instead of relying on the rather naive ‘steady state’
assumption that dung decay and arrival rates are balanced evenly at all survey sites.
Estimating Absolute Densities of Prey from Dung Counts
Theoretically, if we possess good estimates of daily defecation rates (number of dung piles
deposited by an animal/day), we can take the dung survey analyses one step further. By
dividing the daily dung deposition rate (corrected for decay) by the daily defecation rate per
animal, we can estimate the number of animals that produced the dung. Since we know the
total area of the dung plots, we can then estimate absolute densities of prey species.
For example, at both the sites mentioned in the above example we have dung deposition rates
of 5 piles/hectare/day for chital. Assuming a defecation rate of 10 piles/day/chital, we can
estimate the density of deer as 0.5 chital/hectare (50 chital/km2) at these sites.
However, defecation rates of wild ungulates are notoriously variable in nature and very difficult
to measure accurately. Therefore, unless dung-based methods are strongly warranted by local
80
Field Surveys: Assessing Relative Abundances of Tigers and Prey
circumstances and prior knowledge, we prefer the line transect method described in Chapters
9 and 10 for estimating absolute densities of tiger prey.
REFERENCES
Carbone, C., Christie, S., Coulson, T., Franklin, N., Ginsberg, J., Griffiths, M., Holden, J.,
Kawanishi, K., Kinnaird, M., Laidlaw, R., Lynam, A., Macdonald, D.W., Martyr,
D., McDougal, C., Nath, L., Obrien, T., Seidensticker, J., Smith, D., Sunquist, M.,
Tilson, R. and Wan Shahruddin W.N. 2001. The use of photographic rates to estimate
densities of tigers and other cryptic mammals. Animal Conservation 4:75-79.
Ernest, H.B., Penedo, M.C.T., May, B.P., Syvanen, M. and Boyce W.M. 2000. Molecular
tracking of mountain lions in the Yosemite Valley region in California: Genetic analysis
using microsatellites and faecal DNA. Molecular Ecology 9:433-441.
Hiby, L. and Lovell P. 1991. DUNGSURV – a program for estimating elephant density from
dung density without assuming “steady state”. Pages 73-80 in Proceedings of the
workshop for censusing elephants in forests (Eds: U. Ramakrishnan, J.A. Santosh,
and R. Sukumar). Asian Elephant Conservation Centre, Bangalore, India.
Jenelle, C.S., Runge, M.C. and Mackenzie, D.I. 2002. The use of photographic rates to
estimate densities of tigers and other cryptic animals: A comment on misleading
conclusions. Animal Conservation (In press).
Karanth, K.U. and Nichols, J.D. 2000. Ecological status and conservation of tigers in India.
Final Technical Report to the US Fish and Wildlife Service (Division of International
Conservation), Washington, DC, and Wildlife Conservation Society, New York. Centre
for Wildlife Studies, Bangalore, India.
Marques F.F.C., Buckland, S.T., Goffin, D., Dixon, C.E., Borchers, D.L., Mayle, B.A. and
Peace, A.D. 2001. Estimating deer abundance from line transect surveys of dung:
Sika deer in southern Scotland. Journal of Applied Ecology 38:349-363.
Rodgers, W.A. 1991. Techniques for wildlife census in India: A Field Manual. Wildlife
Institute of India, Dehradun, India.
Smith J.L.D., McDougal C. and Miquelle, D. 1989. Scent marking in free ranging tigers,
Panthera tigris. Animal Behavior 37:1-10.
81
 Field Surveys: Assessing Relative Abundances of Tigers and Prey Field Surveys: Assessing Relative Abundances of Tigers and Prey

Thompson, S.K. 1992. Sampling. Wiley, New York, NY, USA. Table 8.1: A Count-based Index of Tiger Density from Five Reserves in India.

Thompson, W.L., White, G.C. and Gowan, C. 1998. Monitoring vertebrate populations.
Academic Press, New York, NY, USA.
Total distance Estimated Index
Shenk, T.M., White, G.C. and Burnham, K.P. 1998. Sampling variance effects on detecting Site
walked (km) n I SE
density dependence from temporal trends in natural populations. Ecological
Monographs 86:445-463.
Kanha 770.7 197 2.61 0.24
Wasser, S.K., Houston, C.S., Koehler, G.M., Cadd, G.G. and Fain, S.R. 1997. Techniques
for application of faecal DNA methods to field studies of Ursids. Molecular Ecology
Kaziranga 631.1 173 2.64 0.28
6:1091-1097.

Yoccoz, N.G., Nichols, J.D. and Boulinier, T. 2001. Monitoring of biological diversity in Nagarahole 1098.0 129 1.17 0.15
space and time. Trends in Ecology and Evolution 16:446-453.
Namdapha 363.1 7 0.19 0.08

Pench 512.0 34 0.67 0.14

(Source: K.U. Karanth unpublished data)

n = Total number of tiger scats encountered

I = Mean number of scats encountered/10 km walked

82 83
 Field Surveys: Assessing Relative Abundances of Tigers and Prey Field Surveys: Assessing Relative Abundances of Tigers and Prey

Table 8.2 A Frequency-based Index of Tiger Density from Five Reserves in India. Table 8.3. A Photo Trapping Rate-based Index of Tiger Density from Seven Reserves in
India.

Total distance Estimated Index Sampling Efforts Index of Tiger Density

Site Site
walked (km) n I SE (Trap nights) n I

Kanha 770.7 66 0.86 0.04 Bandipur 946 31 3.3

Kaziranga 631.1 52 0.83 0.05 Bhadra 587 29 4.9

Nagarahole 1098.0 72 0.66 0.05 Kanha 803 87 10.8

Namdapha 363.1 6 0.17 0.06 Kaziranga 544 80 14.7

Pench 512.0 21 0.41 0.07 Nagarahole 868 52 6.1

(Source: K.U. Karanth unpublished data)

Pench 787 42 5.3

n = Number of segments in which scats were found

Ranthambore 840 73 8.7
I = Mean proportion of 10 km segments in which tiger scats were found
(Source: K.U. Karanth unpublished data)

n = Total number of tiger photos

I = Mean number of tiger photos/100 trap nights

84 85
 CHAPTER 9

STATISTICAL CONCEPTS: ESTIMATING ABSOLUTE

DENSITIES OF PREY SPECIES USING LINE TRANSECT
SAMPLING

Len Thomas1 and K. Ullas Karanth2

Centre for Research into Ecological and Environmental Modelling (CREEM)
1

Mathematical Institute, University of St. Andrews, St. Andrews Fife, Scotland KY 16 9SS, UK.
Email: <len@mcs.st-and.ac.uk>

Wildlife Conservation Society (International Programs), Bronx, New York, NY-10460-1099, USA.
2

Email: <ukaranth@wcs.org>

INTRODUCTION

BLANK Line transect sampling is one among a family of abundance estimation approaches collectively
known as distance sampling methods. These methods are described in detail in Buckland et
al. (2001), which is an update of Burnham et al. (1980) and Buckland et al. (1993). In
addition, a book containing details of the latest research in this area is forthcoming (Buckland
et al. in prep.). In this chapter, we provide a basic “crash course” in line transect sampling for
estimating density and abundance of tiger prey species. Users are referred to the above texts
for more in-depth information.

In a line transect survey an observer traverses a series of transect lines, counting any animal
that he or she visually detects (Figure 9.1). Additionally, the observer records the perpendicular
distance from the line to the animal (or, if that is not possible, the radial distance and sighting
angle) in each case (Figure 9.2).

To understand the line transect method in the context of the overall sampling framework
mentioned in Chapter 3, we can start by considering a related method called the strip transect.
In a strip transect survey the observer traverses a series of lines, counting all animals of the
target species that occur within a fixed distance, w, of the line. We can then use the canonical
estimator from Chapter 3 to estimate the number of animals in the survey region. Recall
equation 3.4 from Chapter 3,

C′
Nˆ =
pˆ α

where Nˆ is estimated abundance, C ′ is the number of animals counted, pˆ is the estimated

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 Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
proportion of animals counted and α is the proportion of the study area that was surveyed.
pˆ relates to observability, while α relates to spatial sampling. In strip transects, we assume
that all animals in the surveyed region were counted, so pˆ = 1. To calculate α , we can think
of the area surveyed as a long, thin rectangle of width 2w and length L, where L is the total
length of the lines. The area surveyed is therefore 2wL, and the proportion of the total survey
area, A, surveyed is α =2wL/A. Hence,
C′ AC '
Nˆ = =
pˆ (2 wL A) 2 wLpˆ
The main problem with strip transects is the assumption that all animals in the surveyed
region are counted. For this to be realistic, the strip must be narrow, so that we can be sure we
do not miss anything within it. However, when the strip is narrow we will inevitably see many
animals outside the strip, and these must be ignored. Such wastefulness with data may be
acceptable for very common species, but is frustrating and inefficient for scarce species. For
this reason, strip transects are sometimes used for dung surveys of tiger prey (Chapter 8), but
are almost never used for surveys of the animals themselves.
The line transect method is a generalization of the strip transect. In line transects we do not
have to assume that all animals are counted. Instead, we record additional information that
allows us to estimate the proportion of animals counted. This means we can use a much wider
strip for recording animals – indeed in most line transect studies of tiger prey we record all
animals of the target species that we see.
In a line transect survey, the observer traverses a series of lines, and for each animal detected,
the observer measures the distance from the line to the animal. For tiger prey studies, we need
to know what species the animal is, so we count “detection” only if the animal is seen (rather
than just heard). Not all animals that the observer passes are detected (Figure 9.1), but a
fundamental assumption of the basic method is that all animals that are actually on the line
are detected. Intuitively, we would expect that animals become harder to detect with increasing
distance from the line – animals farther from the line are more likely to be hidden by vegetation,
and are more cryptic, etc., so we are more likely to miss them as we pass.
This means that if we plot a histogram of number of detections against distance from the line,
we expect smaller numbers of detections with increasing distance (Figure 9.3). The key to
distance sampling analyses is to fit a detection function to the observed distances, and to use
this fitted function to estimate the proportion detected, pˆ . Once we have an estimate of pˆ ,
we can use the canonical estimator, just as with the strip transect method, to estimate the
number of animals in the survey region.
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Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
Line transect methods, and other types of distance sampling surveys, have been used
successfully in a very diverse array of taxa, including trees, shrubs and herbs, insects,
amphibians, reptiles, birds, fish and marine and terrestrial mammals. Although they can
potentially produce very reliable estimates of abundance, it is crucial to be aware of the
method’s assumptions (see below) and to use appropriate field techniques to ensure that these
assumptions are met.
Line transect data are commonly analyzed using the free PC software DISTANCE (Thomas
et al. 1998, 2001). At the time of writing, the most recent version is DISTANCE 3.5, which
is the first version to feature an easy-to-use windows-based interface. A prototype of
DISTANCE 4.0 is also available – this version also includes a built-in GIS and a survey
design tool.
LINE TRANSECT THEORY
In this section, we outline how density and variances are estimated in line transect sampling.
It is not essential to understand this material on first reading; for a more complete treatment,
with longer explanations, see Buckland et al. (1993, 2001).
Estimation
In a line transect survey, k lines of length l1 ... lk (with total length L) are positioned within the
study region of area A, according to some randomized design. An observer or team of observers
moves along the line, carefully scanning for prey species of interest. In theory, animals farther
than w from the line are ignored, making the surveyed area 2wL. In practice, for most surveys
of tiger prey, all animals seen on the survey are recorded, and w is set at the data analysis
stage (see Data Analysis, below).
For each animal detected, the observers measure the perpendicular distance, x, from the line
to the animal (some animals occur in groups or clusters – these are dealt with below). In
practice, observers usually measure the actual distance to the animal, r, and the angle of the
animal from the line, θ . These are converted to perpendicular distances using x = r sin θ
(Figure 9.2).
Suppose that C ′ animals are detected on a survey. Let p be the probability that a
randomly chosen animal within the surveyed area is detected, and suppose that an estimate
of this, pˆ , is available. As we saw in the Introduction, animal abundance, N, is then
estimated by
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 Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling

AC ′ distance, x, from the line to the centre of the cluster, and the number of animals in the
Nˆ = cluster, s.
2 wLpˆ

Animal density, D, is abundance divided by area, and so is estimated by The above formulae then give the density of clusters in the study area. To obtain the density
of individuals, we must multiply by the expected cluster size, E(s):
C′
Dˆ =
2 wL pˆ C ′fˆ (0) Eˆ ( s )
Dˆ =
2L
The problem now is to estimate p. We start by defining the detection function, g(x), to be the
probability that an object at distance x from the line is detected, given that x is between 0 and If large clusters and small clusters are equally visible, then we can estimate E(s) as the mean
w. We assume that g ( 0) = 1 , i.e., we are certain to detect an animal on the trackline. We then of the observed cluster sizes. However, this is rarely the case. More commonly, large clusters
w
define a new parameter, µ , as the area under the detection function, ∫ g ( x ) dx . µ is called are more visible, which means that large clusters are over-represented in the sample. This is
0
the effective strip (half-) width, and can be thought of as the distance at which as many called size bias. There are several ways of estimating E(s) in the presence of size bias (Buckland
objects are detected beyond µ as are missed within µ (Figure 9.4). p is related to µ et al. 1993, 2001). One that works well in practice is to regress log s on gˆ ( x ) , the estimated
because p = µ w . Therefore, probability of detection ignoring cluster size, and then to predict log s where detection is
certain, gˆ ( x ) = 1 , as there can be no size bias where all clusters are seen. The prediction is
C′ C′ C′ then back transformed, using a bias adjustment. This is the default method used in DISTANCE,
Dˆ = = = but other options are available.
2 wL pˆ 2 wL ⋅ µˆ w 2 µˆ L

We now need an estimate, µˆ , of µ . To do this, we use another function, called the probability Variance and Interval Estimation
density function (pdf) of perpendicular distances, f(x). If the density of animals perpendicular The variance of Dˆ is well approximated using the formula
to the track line is constant, then this function is simply the detection function g(x) rescaled so
that the area underneath it is 1 – i.e., f ( x ) = g ( x ) / µ . In particular, because we assume that
g ( 0) = 1 , it follows that f (0) = 1 / µ (Figure 9.4). Hence, vaˆr( D ) = D 2  +
( )+
( )
 vaˆr (C ′) vaˆr fˆ (0) vaˆr Eˆ ( s ) 

 C ′ fˆ (0) 2 Eˆ ( s ) 2 
2

C ′ C ′fˆ (0)
Dˆ = = The variance of the numbers of animals (or clusters) seen var(C ′) is generally estimated
2 µˆ L 2L
from the variation in counts among lines (after accounting for differences in line length). For
The problem is now reduced to modelling the pdf of perpendicular distances, and evaluating reliable estimation of this variance, at least 15-20 lines are needed. If there are fewer lines,
the fitted function at x = 0 . Fitting a density function is a standard statistical problem, and then a value for var(C ′) can be estimated by assuming that vaˆr(C ′) is proportional to E (C ′) .
there is a large literature available to us on the topic. DISTANCE uses a flexible and robust ( )
A likelihood-based estimate of var fˆ (0) is a by-product of the detection function estimation,
method in which a parametric “key” function is selected and, if it fails to provide an adequate ( )
and the method of estimating var Eˆ ( s ) depends on the size bias methods used. The software
fit, polynomial or cosine series adjustments are added until the fit is judged to be satisfactory DISTANCE can calculate all of these variances automatically.
by one or more criteria (see Data Analysis, below).
A useful quantity to calculate is the coefficient of variation of the estimate,
Clusters of Animals
0 .5
Tiger prey species often live in groups. Therefore, in line transect surveys we often encounter  vaˆr( Dˆ ) 
Cˆ V ( Dˆ ) =  
 Dˆ 
2
groups of animals rather than solitary individuals. In this case, we treat the group, or cluster,
as the object of interest. For each cluster seen, we measure the perpendicular

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 Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
This gives a measure of the size of the variance, relative to the size of the estimate. Because
it is unit-less, CVs can be compared between studies, to see how precise the study is. For a
study of tiger prey, a CV of 15% or less would be considered good.
If we assume that Dˆ is log-normally distributed, approximate 95% confidence limits are
given by Dˆ C , Dˆ ⋅ C , where
{ [
C = exp 1.96 vaˆr(ln Dˆ ) ] }
0.5
with

(
vaˆr(ln Dˆ ) = ln 1 + Cˆ V ( Dˆ ) 
2
)

Generally, we prefer to use bootstrap variance and interval estimates for the final estimation.
Bootstrapping is a non-parametric statistical method, which avoids the need to make any
assumptions about the distribution of the estimated quantities. In bootstrapping, multiple
random resamples are taken from the data, and for each resample, Dˆ is calculated. The
variance of the resampled Dˆ s is taken as an estimate of var(Dˆ ) . Because each transect line
is considered to be independent, resamples are taken at the level of the line, not the observation
(see Buckland et al. 1993, 2001 for more details).
ASSUMPTIONS OF LINE TRANSECT SAMPLING
There are four critical assumptions in line transect sampling, which must be met for reliable
estimation of density or abundance.
Assumption 1. Transect lines are located randomly with respect to the distribution of
animals.
This assumption is used at two levels. Firstly, it allows us to use the observed distribution of
perpendicular distances to estimate the detection function, and hence average probability of
detection, p. Secondly, it allows us to extrapolate from what is observed during the survey to
the study area as a whole. If this assumption is not met, then the estimates will be biased. For
example, if transect lines are located along existing trails, and these trails are also used by the
animals, then we will overestimate abundance.
The best way to ensure that this assumption is met is to place the lines using a random
or systematic design (see Survey Design, below). It is then not necessary to make
any assumption about the distribution of the animals themselves (e.g., that they are
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Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
distributed randomly, uniformly, or clumped, etc.) since our inferences will be valid
regardless of the underlying pattern of spatial distribution of animals.
If the same transect lines are used repeatedly, then there is a danger that animals will either
start using the lines as paths or, in heavily hunted areas, that they may be repelled by the
perceived danger from humans. Therefore, in forested habitats, it is very important to do as
little cutting of lines as possible – just enough to enable you to move quietly along the line
(see Chapter 10). If there is a suspicion that animals are reacting to the transect lines, then it
is worth doing a separate study involving strip transects of animal dung at fixed distances
from the worst affected lines to determine the extent of the problem.
Assumption 2. Animals on the line are detected with certainty.
As we saw in the previous section, a fundamental part of the derivation of the density estimator
was the assumption that all objects at zero perpendicular distance are detected – i.e., that g(0)
= 1. If we miss objects on the trackline, then we will underestimate density. Bias is a simple
function of g(0) – for example if we miss 10% of the animals on the line then our density
estimate will be on average 10% too low.
For some line transect studies (e.g., shipboard surveys of whales, which spend a significant
proportion of time below the water), this assumption is very difficult to meet. Advanced
methods, outside the scope of this chapter, must be used to estimate g(0). For studies of
ungulate prey species, it should be possible to ensure that g(0) = 1 through good field methods,
as outlined in Chapter 10.
Assumption 3. Animals are detected at their initial location.
This assumption is easy to satisfy for studies of stationary objects (e.g., plants or animal
dung), or where the observer is moving at high speed (e.g., aerial surveys). However, for
surveys of tiger prey, where the observer is on foot, this can be a major concern.
Studies have shown that movement does not cause a serious problem provided that: (i)
movement is “random” – i.e., not in response to the observer; and (ii) the observer is moving
substantially faster than the animals (at least 3-4 times faster, Buckland et al. 1993, 2001).
These conditions can conceivably be met if the target animals are grazing (in which case they
will not be moving fast), provided that the observer sees them before they become aware of
the observer’s presence.
A bigger problem for studies of tiger prey is evasive movement prior to detection. This occurs
when animals sense the approaching observer, and move away from him or her before the
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 Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
observer detects the animal. When the observer finally sees the animal, its position is farther
from the line than it “should” be. This results in too few detections of animals close to the
track line, and too many a short distance away from the line (Figure 9.5).
Methods of dealing with this situation during analysis are discussed later, but the important
thing is to avoid it happening in the first place.
Therefore, it is vital that observers move as silently as possible along the transect lines, and
concentrate their effort on detecting animals in the region of the line but far enough ahead of
them that they have not yet detected the observer. The ideal is to spot a herd of chital grazing,
to stop and carefully measure the distance and angle to the center of the group, carefully
count the number of animals in the group, and only then allow the animals to detect you.
Some prey species, such as Hanuman langur are simply too wary for this assumption to be
met for foot-based observers, and some studies have used elephant-back surveys instead
(Karanth 1988).
Evasive movement prior to detection in, say, 5% of observations will cause no problems for
the analysis. If, however, it happens in a substantial proportion of observations, then it is
likely that density will be underestimated.
Assumption 4. Measurements are exact.
Ideally, the recorded distances and angles are exact, without measurement errors,
recording errors or rounding. Although the effect of inaccurate measurements can often be
reduced by careful analysis, it is better to gather good data in the field, rather than to rely on
analytical methods. Good field practices are described in Chapter 10.
One problem that sometimes occurs in surveys of tiger prey is rounding of angles
close to the transect line. This happens when animals or clusters of animals
are seen a long way ahead, in the vicinity of the transect line. Instead of carefully
measuring the compass bearing of the animal or center of the cluster, the observer erroneously
writes down the same compass bearing as the transect line. This translates into a
perpendicular distance of zero, when the true distance could be up to 10 or 20 meters
away. When this happens, the data show a “spike” at zero distance (Figure 9.5), leading to
problems at the analysis stage. This problem can be completely avoided by proper field
methods.
Other Assumptions
Some other assumptions are often mentioned in the literature, but they are not nearly
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Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
as important for reliable estimation of density or abundance. One is that detections should
be independent events. If detections are somewhat dependent (e.g., detecting one animal
leads to it fleeing and frightening others, which are then detected), then the theoretical
variances will be underestimated. However, we recommend the use of robust empirical estimates
of variance, based on bootstrapping, which only assumes that detections are independent
between transect lines. There is a minor issue to do with model selection of the detection
function, but this has little practical implications. An extreme form of non-independence is of
animals that occur in groups, and we have outlined methods for dealing with them in a
previous section.
Another issue that is sometimes mentioned is that animals should not be counted more than
once on the same line. This is only an issue if it occurs commonly, or if there is a possibility
of “chasing” an animal down the line, counting it many times. It is not a problem if the same
animal is counted from different lines, again, so long as it has not been “chased” from one line
to the other.
We do not need to assume that all animals at the same distance from the transect line, have
the same probability of detection. Clearly, the probability of detection will be different
for each animal, depending on the observer, habitat, individual animal behavior, and a
host of other factors. We are only interested in estimating p, the average probability
of detection for the species during our survey. We use distance from the line to enable us to do
this because, given assumption 1 above, we know the true distribution of animals with respect
to the line. We can try to improve the precision of our estimate of p by including other factors
that may be important to the detection process – this is called multiple covariate distance
sampling (Thomas et al. 2001), and is beyond the scope of this chapter.
Lastly, there is one important practical consideration for reliable estimation of the detection
function, relating to the shape of the detection function near zero distance. Distance
sampling theory performs well when a “shoulder” in detectability exists near the line – i.e.,
where detectability is certain near the line and stays certain or nearly certain for some distance
(Figure 9.6). If detectability falls sharply just off the line or point, then estimation tends to be
poor, and the analysis can be unreliable. This is not an assumption, but a property that allows
more robust estimation, and is often referred to as the shape criterion.
SURVEY DESIGN
A survey design is a protocol for placing transects within in the study area. Designing a
survey involves making three (often linked) decisions: (i) what length, orientation, and shape
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 Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling

of transect to use (called the sampler geometry); (ii) how to place the transects in the study the density or abundance estimate will be. Covering the same line many times in a short time
area; and (iii) what sample size of transects to use. These are covered briefly below, but are period does not constitute replication in this sense, and so is much less useful than lines
discussed in much greater detail in Chapter 7 of Buckland et al. (2001), and in a chapter of placed in new locations. If the lines are being laid out with long-term monitoring of trends in
Buckland et al. (in prep.). The new software, DISTANCE 4.0, has a built-in survey design mind, then it is better to survey the same set of lines on each occasion (see Skalski 1990, for
module, which allows many of these options to be explored using a digitized map of your a discussion of survey design for monitoring).
study area.
Some form of randomization should be used in placing the transect lines, otherwise the
Sampler Geometry observations on the lines cannot validly be extrapolated to the study area as a whole. While it
is acceptable to locate the transect lines completely at random within the study area, a better
In most studies of tiger prey species, observers walk along transects in the early morning or
design is a systematic grid of transects with random starting point (Figure 9.7). Systematic
late afternoon, when most of the species are most active and so the number of sightings is
designs give a more even spatial coverage of transects, which results in lower variance.
maximized. The length of the transect is thus set by the distance that an observer can cover
(Advanced readers may notice that we have ignored the possibility of model-based estimation
during a 3-4 hour observation period – typically 3-4 kilometers.
of density, which can be used to get density estimates from lines that are subjectively located.
However, such estimates can easily be biased, and the analysis requires a black-belt in statistics
In many studies, transects are continuous straight lines. One problem is that time must be
to perform properly. For more information about model-based sampling, see Thompson (1992).
spent getting to the start of the line, and returning from the end. A more convenient alternative
is to use a continuous shape, such as a square or equilateral triangle (Figure 9.7). The survey
The design can sometimes be improved further using stratification. This is where the study
would start from the most convenient point (e.g., the point closest to a road, or where the line
area is divided into different regions, called strata, and each region is treated independently.
crosses a road), and move around in either direction (one could alternate directions) until the
One reason to do this is when we want different estimates of abundance in the different strata.
observer returns to the starting point. The sides of the sampler are treated as one continuous
Another reason is to improve precision of the overall estimate of abundance, by defining
line for the purposes of analysis. Curved transect lines could also be used (Hiby and Krishna
strata that are expected to differ in animal abundance – for example, different habitats (forest
2001), but care must be taken to avoid bias by subjective placement.
and grassland). As a general rule, for minimum variance, we want to construct strata so that
transects within strata are as similar as possible, and those between strata are as different as
In many cases, the orientation of transects is not important, and can be chosen at random.
possible. Another possible way to decrease variance is to allocate more transect lines per unit
However, where there is a known gradient in density, and the transect is a straight line (rather
area to strata expected to contain a greater abundance of animals. However, this is not usually
than a square or other shape), it is best to orient it, so that one end of the transect tends to
practical in multi-species surveys, and it is often better (and simpler) to keep the same density
sample the low-density area and the other end samples the high-density area. For example, if
of transects in all strata. In this case, one can ignore stratification completely at the design
transects are located in a mountainous area they should be oriented up and down the slope if
stage, and use post-stratification during analysis (see Buckland et al. 1993, 2001 for more on
possible. This reduces the variance of the density estimate – a general rule of thumb for
this). Avoid creating too many strata – remember that each needs at least 15-20 transects for
minimum variance is that each transect should contain as much of the variation in density as
reliable estimation of variance.
possible, so that the between-transect variation is minimized. Another way to reduce variance
is through stratification (see below).
Sample Size
Placement of Transects How much sampling effort is required in a particular study? Firstly, a minimum number of
observations are required in order to be able to reliably model the detection function. The
Three main concepts should guide the placement of transect lines within the study area:
number required depends on the characteristics of the data – for example data showing any
1) replication; 2) randomization; and 3) stratification.
tendency to a spike (Figure 9.5) requires a much greater sample size than “good” data with a
wide shoulder. As a rule of thumb, it is often difficult to get a robust result with fewer than
Spatial replication of lines is important for reliable estimation of variance. As a minimum, at
60-80 observations, although this is very variable. Remember that for animals that occur in
least 15-20 lines are required. After this, the more lines you have, the smaller the variance of

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 Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
clusters, this means 60-80 observations of clusters. As we have said before, we also require
at least 15-20 transects to reliably estimate variation in numbers seen between lines.
Beyond these minima, the answer depends upon the required precision of the estimates. Recall
that the variance of density is estimated by:
vaˆr( Dˆ ) = Dˆ 2  +
( ) +
( )
 vaˆr (C ′) vaˆr fˆ (0) vaˆr Eˆ ( s ) 

 C ′ fˆ (0) 2 Eˆ ( s ) 2 
2
This shows that there are three separate components to the variance – variation in number
counted, var(C ′) , variation in estimating the detection function (or more properly f(0))
and variation in estimating expected cluster size, E(s) (for animals in clusters). More sampling
effort will generate more observations for estimating f(0) and E(s), so the variance of
these components will decrease. The situation is not so simple for var(C ′) . This is best
estimated using variation in counts between lines. Therefore sampling more on the same lines
(e.g., by repeating the same lines, or extending them) tells us little about the variation
in counts over space. So, it is almost always better to sample new lines if extra effort is
available. The exception is when establishing a new line is very much more costly than
surveying the same line again. Formulae for calculating the expected CV and variance of
estimates for a given level of sampling effort, from pilot survey data, are given in Buckland
et al. (1993, 2001).
DATA ANALYSIS
Data Entry and Recording
Data are usually entered in the field onto a data form such as that shown in Appendix 3.0.
The observer must ensure that all data are accurately recorded in the field. On returning to the
field station, the observer should verify the data. Data should be entered into the computer as
soon as possible, typically using spreadsheet or database software (e.g., MS-EXCEL,
MS-ACCESS, LOTUS, etc.). This enables further checking and correction of errors
at an early stage. Data recording and entry errors can cause intractable problems in later
analyses.
Preliminary Checks
We strongly recommend the use of the software DISTANCE, versions 3.5 or 4, for analyses
of line transect data. However, during initial training and exposure to line transect surveys,
we have found it to be very useful for the trainees to calculate parameters by hand. One
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Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
invaluable exercise is for trainees to plot a histogram of their detection distances, and check
it for evidence of problems as outlined previously. A curve can be fit to the histogram by eye,
and an estimate of the probability of detection can be made as in Figure 9.3. From this, a
preliminary estimate of density can easily be calculated. Such rough point estimates can be
later compared to the more sophisticated estimates generated by DISTANCE.
Data Analysis Using DISTANCE
The procedures for the final analyses of line transect data are explained in detail by Buckland
et al. (1993, 2001), and in the manual that accompanies the program DISTANCE (Thomas
et al. 1998, 2001). Here, we provide just a brief overview of the steps involved. We have tried
to steer clear of a cookbook approach to analysis and, above all, recommend that you do not
unthinkingly use the DISTANCE defaults.
First, the transect data must be entered into a DISTANCE database file, called a project file.
Although the field data can be manually keyed into DISTANCE, it is simpler and less error-
prone to transfer the data from a spreadsheet or database application using DISTANCE’s
import facility.
Each analysis in DISTANCE has two components, a data filter and a model definition. The
data filter is used to select a subset of the data – for example only data for one species. Data
filters are also used to truncate data (see below). The model definition specifies how these
data should be analyzed – for example what model to use for the detection function, and how
variances should be calculated.
Four main steps are involved in a complete analysis of line transect data:
1. Data exploration and truncation
2. Selection of an appropriate model or models for the detection function
3. For species that occur in clusters, estimating the expected cluster size
4. Deriving estimates of density and other parameters using the selected model
Data Exploration
It is difficult to do a good analysis of a line transect survey without thoroughly understanding
the data first. So, the first stage of any analysis is exploratory. We recommend a visual
approach, where DISTANCE is used to plot histograms of the data, splitting the observed
distances into a large number of intervals (15-20), particularly near the origin (at x = 0; e.g.,
Figure 9.6). Look for evidence of a shoulder in the histograms. Check the plots for evidence
of violation of the assumptions: A spike at 0 distance, signs of evasive movement and
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 Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
“rounding” or “heaping” of distance data at specific distances. A simple detection function
model, such as the half-normal or hazard rate (see below) can be fit to the data, and the
goodness-of-fit statistics used to judge whether any violations of the assumptions are important
or not. Guidelines for dealing with problematic data are given in the texts referred
to above.
Part of the exploratory process is deciding where to truncate the data. It is common to discard
around 5-15 % of the observations with the largest distances before proceeding to serious
model selection. These data are usually part of a long “tail” of observations, which are of no
help in choosing among the different possible detection functions, and indeed tend to encourage
“over-fitting” (see next section).
Modeling the Detection Function
DISTANCE uses a flexible approach to fit a model of the detection function to the observed
data. To start with, a key function is fit to the data. Functions available are uniform, half-
normal and hazard rate (there is also a negative exponential function, but we do not recommend
its use). The fit can then be improved by introducing some adjustment terms into the model.
These terms modify the shape of the key function, to help it fit the data better. Adjustment
terms can be either polynomial or cosine series expansions. (This is covered in much greater
detail in Buckland et al. 1993, 2001).
The basic goal is to produce a plausible model for the detection function using as few parameters
as possible. The more parameters that are used, the more closely the estimated function will
fit the observed data, but the less likely that the model will generalize to other data sets that
could have been collected under the same circumstances. Fitting too many parameters is
called “over-fitting”.
One way to objectively determine which model to use is via the Akaike Information
Criterion (AIC) statistic. For well-behaved data, which do not appear to violate any of the
basic assumptions, we recommend the following procedure. Choose a small number
of candidate key function-adjustment term combinations (see Buckland et al. 1993, 2001
for recommended combinations). For each, use the automated forward-selection
procedure in DISTANCE to select the number of adjustment terms that gives the lowest AIC.
Then, compare these models and, unless you have some doubts about the fit of any of them,
choose the one with the lowest AIC. If two models have very similar AICs (less than 1
difference) then it is difficult to choose among them. This “model selection uncertainty”
can be accounted for in the final density estimate, using bootstrapping (Buckland et al.
1993, 2001).
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Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
It is vital to realize that the above procedure will not produce a good estimate if there are any
problems in the data, such as rounding or evasive movement – if the data show any violations
of the key assumptions then the analysis may be salvaged using methods outlined in Buckland
et al. (1993, 2001). Possible measures include “binning” the data into intervals before analysis,
truncating more data, subjectively choosing detection function models, and performing
additional studies to estimate the extent of any bias. The best approaches depend on the cause
and scale of the problem.
Cluster Size and Size Bias
The ecological propensity of different prey species to occur in clusters varies greatly. Solitary
species such as muntjac or sambar are usually encountered in small clusters of 1-3 animals.
Social species such as chital, barasingha or gaur can occur in clusters ranging in size from
one to several dozens of animals.
As we discussed in the section on Line Transect Theory, when species occur in clusters
of animals, we treat the cluster as the unit of observation and record the cluster size separately.
The detection function gives the probability of observing a cluster of animals, and we
must independently determine the expected cluster size. It is quite common for probability
of detection to be a function of cluster size (“size bias”): Large clusters of animals are
easier to see. This leads to an overabundance of large clusters in our observations, so
that the mean of the observed clusters is an overestimate of the mean cluster size in the
population.
There are many potential ways of dealing with this size bias (Buckland et al. 1993, 2001).
One is to include cluster size as another variable in the detection function model, in addition
to perpendicular distance. Such an option is available in DISTANCE 4. Another, well proven
method, which is the default in DISTANCE 3.5, has been outlined in the section on Line
Transect Theory.
Parameter Estimates and Measures of Precision
Once the modeling is finished, DISTANCE can be used to derive estimates of density and
abundance, and of variance, confidence intervals and CVs, using the methods outlined in the
Line Transect Theory section. So long as there are a good number of transects (at least 20),
then bootstrapping can be used to derive more robust estimates of variance and confidence
intervals.
Once the final estimates have been obtained, it is important to consider how reliable they may
be. The importance of any possible violations of the key assumptions must be considered
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 Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling

and, if necessary, extra fieldwork or computer simulations performed to determine the extent 4.0. Beta 1. Research Unit for Wildlife Population Assessment, University of St.
of any biases. With good field methods, and a well-informed analysis, line transect methods Andrews, Scotland, UK.
should produce very reliable and precise estimates of density and abundance for all the principal
prey species of the tiger. Thompson, S.K. 1992. Sampling. Wiley, New York, NY, USA.

REFERENCES

Buckland, S.T., Anderson, D.R., Burnham, K.P. and Laake, J.L. 1993. Distance Sampling:
Estimating abundance of biological populations. Chapman and Hall, London, UK.
Reprinted 1999 by Research Unit for Wildlife Population Assessment, University of
St. Andrews, Scotland, UK.

Buckland, S.T., Anderson, D.R., Burnham, K.P., Laake, J.L., Borchers, D.L. and Thomas,
L. 2001. Introduction to distance sampling. Oxford University Press, Oxford, UK.

Buckland, S.T., Anderson, D.R., Burnham, K.P. and Laake, J.L. Borchers, D.L. and Thomas,
L. (Eds.). Advanced Distance Sampling. Oxford University Press, Oxford, UK. (In
Prep).

Burnham, K.P., Anderson, D.R. and Laake, J.L. (1980). Estimation of density from line
transect sampling of biological populations. Wildlife Monographs 72:1-202.

Hiby, L. and Krishna, M.B. 2001. Line transect sampling from a curving path. Biometrics
57:727-731.

Karanth, K.U. 1988. Population structure, density and biomass of large herbivores in south
Indian tropical forest. M.S. thesis, University of Florida, Gainsville, USA.

Skalski, J.R.A. 1990. Design for long-term status and trends monitoring. Journal of
Environmental Management 30:139-144.

Thomas, L., Derry, J.F., Buckland, S.T., Borchers, D.L., Anderson, D.R., Burnham, K.P.,
Strindberg, S., Hedley, S.L., Burt, M.L., Marques, F.F.C., Pollard J.H. and Fewster,
R.M. 1998. DISTANCE 3.5. Research Unit for Wildlife Population Assessment,
University of St. Andrews, Scotland, UK.

Thomas, L., Laake, J.L., Strindberg, S., Marques, F. F.C., Borchers, D.L., Buckland, S.T.,
Anderson, D.R., Burnham, K.P., Hedley, S.L. and Pollard, J.H. 2001. DISTANCE

102 103
 Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
Figure 9.1. Diagrammatic representation of a transect survey: A number of transect lines
within the study area are ‘walked’, and perpendicular distances recorded to all animal clusters
detected. Note that not all clusters are detected.
Figure 9.2. The perpendicular distance to the centre of the animal cluster is calculated using
the radial distance r and the sighting angle q, which are related to the perpendicular distance
x by the formula x = r sin θ .
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Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
Figure 9.3. Conceptual basis for line transect sampling: (a) the expected number of animals
detected in eight distance classes if no animal were left undetected; (b) real data, where we
can see a strong tendency to detect fewer animals at greater distances from the line; (c) a
smooth function has been fit to these data by eye. The area under the curve represents animals
seen, and the area above the curve represents animals missed. The proportion seen, p, is
estimated from the area under the curve divided by the total area.
(a)
(b)
(c)
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 Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling

Figure 9.4. Method used by DISTANCE to estimate probability of detection, pˆ ; (a) some
real data; (b) a detection function g(x) is defined, where g(0) = 1. µ, the effective strip width,
is the point at which as many animals are seen beyond µ as are missed before µ. pˆ = µˆ / w .
(c) to estimate µ, DISTANCE fits a probability density function (pdf) to the data. A pdf is a
function where the area under the function is 1. To estimate µ, µˆ = 1 / fˆ (0) .
Figure 9.5. Examples of poor line transect data: (a) evasive movement - animals close to the
line have moved away prior to detection (this is not always so obvious in real data); (b) spike
data- probably caused by rounding of perpendicular distances or angles to 0.

(a)
(a)

(b)

(b)

(c)

106 107
 Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling Statistical Concepts: Estimating Absolute Densities of Prey Species Using Line Transect Sampling

Figure 9.6. Example of good line transect data, which meet the shape criterion. The data Figure 9.7. Examples of different sampler geometries for line transect surveys.
close to the track line have been divided into smaller classes. This can be done in DISTANCE,
and is good practice during the exploratory phase of analysis.

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 C H A P T E R 10

FIELD SURVEYS: ESTIMATING ABSOLUTE DENSITIES OF

PREY SPECIES USING LINE TRANSECT SAMPLING

K. Ullas Karanth1, Len Thomas2 and N. Samba Kumar3

Wildlife Conservation Society (International Programs), Bronx, New York, NY-10460-1099, USA.
1

Email: <ukaranth@wcs.org>

Centre for Research into Ecological and Environmental Modelling (CREEM)

2

Mathematical Institute, University of St. Andrews, St. Andrews Fife, Scotland KY 16 9SS, UK.
Email: <len@mcs.st-and.ac.uk>

Centre for Wildlife Studies, 26-2 Aga Abbas Ali road (Apt. 403), Bangalore, Karnataka-560 042, India.
3

Email: <wcsind@bgl.vsnl.net.in>

INTRODUCTION
BLANK Within the sampling-based framework of animal abundance estimation methods presented in
Chapter 3, line transect survey is a powerful approach to the estimation of densities of ungulates
that are the tiger’s principal prey (Karanth and Sunquist 1992; Kumar 2000). However, it is
important to ensure that the key assumptions of line transect sampling (Chapter 9) are
adequately satisfied when field surveys are carried out. Biological and logistical considerations
often impose severe constraints on being able to execute theoretically justifiable line transect
surveys in the field. In the following sections, we examine how you can try to achieve a
balance between the theoretical needs of line transect sampling, and some of the practical
problems that crop up in the field.

We would however, like to caution the readers that as in all other sampling-based
approaches to animal population estimation, sample size considerations play a critical
role in generating reliable estimates of prey densities from line transects. If densities
of any prey species are low in the area surveyed, despite a lot of sampling effort
(long distances walked), the sample size attained (number of animals or clusters of
animals detected) will be small, and the density estimate derived will be relatively
weak. However, it is very likely that tigers too face the same problem as we do in such
areas! They may not frequently be able to find and catch prey species that occur at
relatively low densities. These low-density species, therefore, may not be important items in
tiger diet. However, if densities of all prey species are low, then you can only record their
distribution (Chapters 4, 5, and 6) rather than estimate relative or absolute densities using
line transects.

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 Field Surveys: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
WHAT ARE “TRUE” TRANSECT LINES?
Establishing valid transect lines is the first key step in carrying out surveys. It is also a step
that is most often misunderstood and misapplied in field surveys. Remember from Chapter 9
that transects have to be randomly located with respect to the distribution of animals. Without
such randomness we cannot extrapolate from what we estimate from the transect lines to the
study area as a whole. For example, imagine a survey where transects are located along
existing animal trails within a reserve. It is quite likely that animal density along the trails
will be higher than away from trails, so we will see a greater density of animals on our survey
than we would if the lines were located at random. In addition, the possibly greater number of
animals near the trails will be mistakenly assumed to reflect the shape of the detectability
function rather than a gradient in density. Both factors associated with non-random transect
location will result in over-estimation of animal density.
Now imagine that the survey takes place in an area along trails heavily used by human
hunters. In this case animal density along trails is likely to be lower than away from trails, as
animals will be avoiding potential hunters. The survey personnel will encounter fewer animals,
and will encounter them farther from transects than if the transects were located at random,
and you will tend to under-estimate density and abundance. The only way to guarantee unbiased
results is to locate transects using a random survey design, such as completely random or
systematic placement with a random start (see Chapter 9).
All too often, biologists use existing roads, firebreaks, waterways and other convenient linear
formations as transect lines. In almost all cases, these features represent specific microhabitats
for the tiger’s prey species; prey animals are either attracted to or repelled from them. For
example, roads are likely to be a non-random sample of the survey area’s habitats, topography,
human interference, etc., – all factors that may affect animal density.
In some cases, it is only possible to perform surveys along pre-existing features such as
trails. In this case, if animals are orienting with respect to the transect, then the gradient in
density with distance from the transect is confounded with the gradient in detectability with
distance. So not only is the estimate non-representative, you are also not doing a good job of
estimating the detection function. The only time it makes sense to use such formations is
when you believe that animals are not orienting their movements with respect to them. Then,
in theory, you will only get a valid estimate of animal density in the places you surveyed.
There is still no theoretical basis for extrapolating this estimate to the study area as a whole.
If you want to do this, you must prove that average density in the surveyed area (e.g., along
trails) is the same as that over the whole study area – a difficult task. Some possibilities are:
(i) to perform strip transect counts of animal dung at right angles to the trails to see if dung
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Field Surveys: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
density varies with distance from trail; (ii) compare encounter rates on trails with those from
transects placed at random.
Another consideration with using pre-existing features is that they are often somewhat curved,
although this factor can be adjusted for in the analysis (Hiby and Krishna 2001).
In conclusion, we strongly recommend that the survey personnel actually mark and establish
their own transect lines over the area to be surveyed, applying the sampling and survey
design considerations explained in Chapter 9.
PLACEMENT OF TRANSECTS: PRACTICAL CONSIDERATIONS
The most practical way to sample an area for ungulate prey species is to use a systematic
placement of a transect pattern, but with a randomly chosen starting point. This process can
be visualized as drawing transects on an acetate sheet using some sample survey design (such
as those in Figure 9.7), and then placing the entire sheet on the study area map using a
randomly chosen starting point. Ideally, the more spatially replicated lines in the survey design
there are, the better it is for sampling. However, most tiger habitats are forests with difficult
access, and there are usually high costs and logistical problems associated with having more
and more spatial replicates. The number of spatially replicated transects one can establish
depends on resources (manpower, money) and time available to the survey team.
Ideally, for each prey density estimate we generate there should be at least 15-20 spatially
replicated transect lines (Buckland et al. 1993; Thomas et al. 2001). If your goal is to obtain
one density estimate for the entire area, then these replicate lines should sample the entire
area. On the other hand, if you need separate density estimates for subunits such as forest
ranges or habitat types, then you must have at least 15-20 replicates sampling each subunit of
interest. Smaller numbers of spatial replications may require estimating the variance of
encounter rates theoretically rather than empirically (see Chapter 9 for a fuller discussion).
This is generally considered a less desirable option than generating variance estimates
empirically from your own data (Buckland et al. 1993, 2001).
From a purely statistical sampling perspective, ideally, the survey personnel should select a
specific (randomly chosen, or, pre-determined according to some design) azimuth and simply
walk in that direction, looking for animals. However, in reality, because of dense vegetation
and other obstructions like cliffs or rivers, this ideal is rarely attainable. Even worse, such
true random walks will require literally crashing through the bush and trampling the litter,
generating a great deal of noise. A noisy random transect walk would alert the prey animals
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 Field Surveys: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
from great distance and make them systematically move from their initial location, in response
to the survey process itself. Such a survey would seriously violate a key assumption of
distance sampling that there should be no undetected, responsive movement. Failure of this
assumption can seriously bias the survey results.
Therefore, for estimating prey abundance, the transect lines have to be physically cut and
maintained minimally at least, so that progress of survey personnel along the lines is not
unacceptably noisy. When cutting the line, its width should be kept as narrow as possible:
Just enough to allow passage of survey personnel, but not wide enough to attract or repel the
target species or to artificially inflate the detection probability along the line itself.
To ensure that survey personnel are aware of the exact location of the transect line at all
times, the position of the line should be marked clearly at regular intervals using paint or
flagging. In open grassland habitats it may be possible to maintain lines just by paint marking
without the need to cut down the vegetation.
If prey densities are to be estimated for several species, sometimes their specific distributions
may be restricted to different parts of the study area. In such cases, separate sets of transect
lines may have to be established to count different species. However, distributions of several
prey species usually spatially overlap in most reserves. Therefore, once a system of transects
that samples the entire area is established, it is possible to conduct surveys of several prey
species simultaneously.
If several prey species are being counted during the same survey, walking at a speed of about
1.5 kilometer per hour usually enables efficient detection of animals. Assuming that an observer
can employ about 2-3 hours of concentrated sampling effort in one session, transect lines of
3-4 kilometer length can be covered during each replication. If there are to be 20 lines (spatial
replicates), about 60-80 kilometers of total line length may have to be established.
As seen in Chapter 9, if you walk the same transect lines repeatedly (temporal sampling), you
can improve the precision of the estimates of the effective half- strip width (µ) and cluster
size (s). However, only a large number of true spatial replicate lines can increase the precision
of the estimate of the encounter rate ( C ′ / L ).
If the same transect lines are used repeatedly, then there may be a danger that animals will
either start using the lines as paths or, in heavily hunted areas, that they will be repelled by the
perceived danger from humans. As seen above, this may violate the assumption that transect
lines are random with respect to animal distribution. If animals are attracted to the transect
line then density will be overestimated; if they are repelled it will be underestimated. Therefore,
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Field Surveys: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
in forested habitats, it is very important to do as little cutting of lines as possible – just
enough to enable you to move quietly along the line.
If there is a suspicion that animals are reacting to the transect lines, then it is worth doing a
separate study involving strip transects of animal dung (Chapter 8) at fixed distances from
the affected lines to determine the extent of the problem.
MEETING LINE TRANSECT ASSUMPTIONS IN FIELD SURVEYS
When planning a line transect survey in a new area, always do a pilot survey – otherwise your
first real survey will effectively become a pilot! It is very worthwhile to manually plot a
histogram of the recorded distances (or use DISTANCE for this: Figure 9.6), and check for
problems such as a spike at zero distance, rounding of angles or true distances, or evidence of
evasive movement of animals prior to detection (Figure 9.5).
Please recall (Chapter 9) the four key assumptions that underlie valid line transect surveys:
1. Transect lines are located randomly with respect to the distribution of animals.
2. Animals on the line are detected with certainty.
3. Animals are detected at their initial location.
4. Measurements made are exact.
Field protocols should be focused on ensuring that the above assumptions are met. In this
section, wherever a field protocol prescription relates to meeting crucial assumptions, we
have provided the assumption number in curly braces – e.g., {1} means assumption 1.
The location and alignment of transects must be decided upon in advance, using a proper
survey design (Chapter 9). Thereafter, it is a matter of going to the start point and carefully
marking out the line, taking care not to be deflected from the planned line {1}. Cut vegetation
only where absolutely essential to enable the observers to move quietly along the line {1}. It
is not essential for observers to be able to walk exactly on the line (see below), so you do not
need to hack a path through every small bush! It is, however, essential that the observers
know where the line is {4}, so mark the line at regular and closely spaced intervals.
When surveying along a transect move as carefully and silently as possible, but not too
slowly {3}. Make sure you know exactly where the transect line is at all times {4}. It is
acceptable to move off the transect line by a few meters, for example to get around a thick
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 Field Surveys: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
bush. However, you should remember that the “true line” goes through the bush, and all
measurements should be taken relative to this true line {4}. In addition, it is essential to
ensure that any animal that is actually on the line is detected {2} and important to ensure that
animals close to the line are also seen {shape criterion}. So you must be careful that none
could be hiding in the bush, or just on the other side of it. Scan ahead of you, concentrating on
the area in the vicinity of the line {shape criterion}. The objective is to see animals before
they detect you {3}.
Both active and passive detections should be recorded. Active detection occurs if observers
detect the animal before it becomes aware of them. Passive detections occur if the animal
flushes in response to the observers first. In both these cases the distance and angle
measurements should be taken to the initial position of the animals {3}.
To avoid heaping and rounding errors {4}, the distance and angle measurements should be
accurately read and exactly recorded to the nearest integer value.
When sighting an animal cluster, the measurements usually taken are true distance to the
center of the cluster, r, compass bearing to the cluster, θ1 , and compass bearing of the line,
θ 2 (Figure 9.2). Perpendicular distance, x, can then be calculated during the data entry process,
as x = r sin(θ1 − θ 2 ) . True distances should be measured with a range finder, and compass
bearings measured with a sighting compass. Laser binoculars can take both of these
measurements more accurately, but they are much more expensive. All measurements should
be as accurate as possible {4}, and it is particularly important to avoid recording the same
angle for θ1 and θ 2 , unless the animal really is exactly on the line (extremely rare). Sometimes
a cluster of animals straddles the line – even in this case the center of the cluster is rarely
exactly on the line, so again avoid recording the same angle for θ1 and θ 2 , unless they really
are the same.
Because estimating the density of clusters is the first step, distance measurements should be
taken from the line to the approximate geometric center of the cluster. Remember that a
‘cluster’ in transect surveys refers to an aggregation of individual animals that occurs within
the effectively sampled strip, and, is not always the same as a larger social group. We generally
find that treating all animals within a 30-meter radius as one cluster to be a useful practical
rule in surveys of ungulate and primate prey species that occur in large, widely dispersed
social groupings.
If the animals are in a cluster, take the time to accurately count the animals. Only then should
you move forward along the line. If the animals then flush, and it becomes apparent that the
group is larger than you initially counted, record the extra individuals detected.
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Field Surveys: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
If animals are close to the line but move away before you can get an accurate measurement of
radial distance, it is acceptable to walk along the line to a point opposite to their initial
location and measure the perpendicular distance directly if possible.
Other information that is commonly recorded is your own location along the line (say to the
nearest 50 or 100 meters – this is facilitated by placing distance markers along the line), and
sometimes, simple habitat information. This information can potentially be used for advanced
modeling techniques, the scope of which is beyond this chapter.
Using well-trained, highly motivated survey personnel is vital for collecting high quality
data. When surveying for prey species we recommend that a team of two observers should
walk each line, with the first person concentrating on the line and scanning an arc of 30° on
either side of it. The second observer should concentrate on the area not scanned by the first
observer. They should wear cryptic clothing, and footwear that permits silent passage (Figure
10.1).
The observers should walk at a uniform pace of 1.5 km per hour. This will reduce noise levels
and increase chances of hearing animal cues early, thereby increasing subsequent chances of
sightings. However, the line transect surveys are based on modeling visual detections, and,
therefore, purely auditory detections cannot be used as data. A specimen field data entry form
used in line transect surveys is shown in Appendix 3.0.
Sometimes detections may occur after you have gone past the animal on the side of a transect
line. Such back sightings are not a problem if they are not too many and not systematic. After
a detection, when one observer is measuring and recording data, the second observer should
continue to scan the forest ahead of them and search for animals.
Most field problems of data collection can be resolved by checking the forms immediately
after a field survey for possible data entry errors. The memory of a particular detection fades
rapidly (“was that deer you saw at 8.20 AM behind you on the left or right?”). The individual
observer may not even be available for crosschecking suspect data if this task is postponed to
a later date.
A particular survey may have to be abandoned due to darkness, bad weather - and in the
case of potentially dangerous species like elephants and rhinos - due to the animals literally
blocking your path. In such cases the location where the survey was abandoned and
the actual distance covered should be recorded, so that the sampling effort is recorded
accurately on the data form. This is also useful for resuming the survey at the right point later
if necessary.
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 Field Surveys: Estimating Absolute Densities of Prey Species Using Line Transect Sampling Field Surveys: Estimating Absolute Densities of Prey Species Using Line Transect Sampling

SEASON AND TIMING OF LINE TRANSECT SURVEYS

The season in which surveys are conducted is determined by several factors. If some of the
prey species are locally migratory and leave the study area for a part of the year, then the
investigator has to decide whether he/she wants the densities estimated when these animals
are in the area or out of it.

The footfall noise generated while walking on transects, and, the degree of visibility, are both
important practical considerations in picking the season and timing for the survey. Noisy
underfoot conditions seriously affect survey results as explained earlier, and must be avoided.

Greater visibility results in larger number of detections for a given survey effort: More clusters
of prey are encountered because a wider strip is sampled. As a result, sample sizes are larger.
Additionally, undercounting of cluster sizes is minimized with greater visibility, possibly
reducing size bias in the data. However, (in south Asia at least) the undergrowth is usually
most open (visibility is best) in the dry season when the underfoot conditions are most noisy
because of dry leaf litter. In the wetter seasons the converse is true. Therefore, conducting
line transect surveys in the dry season, but after a few early showers have softened the leaf
litter, is often the best practical approach.

Many of the tiger’s ungulate prey species are active and mobile around dusk and dawn, and
tend to lie up during the hotter parts of the day. Therefore, they are more detectable (a wider
effective strip is surveyed) during dusk and dawn, than during the rest of the day. The shape
of the detection function will also differ by time of day, although the underlying densities do
not change. We have generally found it better to carry out surveys during early mornings and
late afternoons to take advantage of improved detection conditions resulting from animal
activity. If you find that animals are becoming more or less detectable with passage of time,
it may be a good idea to ‘balance’ the survey by temporally replicating it from the opposite
2
ends of the lines.
1
3
Other logistical considerations that may affect the survey design are: The time taken to traverse
the transect line, the distance of lines from the base camp and the mode of transporting survey
personnel to and from the lines.
1. Range Finder
2. Clipboard and Data Form
4 3. Binoculars
REFERENCES
4. Compass
Buckland, S.T., Anderson, D.R., Burnham, K.P. and Laake, J.L. 1993. Distance Sampling:
Figure 10.1 - Survey team walking along a transect line (Inset: Field equipment for line transect surveys). Estimating abundance of biological populations. Chapman and Hall, London, UK.

118 119
 Field Surveys: Estimating Absolute Densities of Prey Species Using Line Transect Sampling
Reprinted in 1999 by Research Unit for Wildlife Population Assessment, University
of St. Andrews, Scotland, UK.
Buckland, S.T., Anderson, D.R., Burnham, K.P., Laake, J.L., Borchers, D.L. and Thomas,
L. 2001. Introduction to distance sampling. Oxford University Press, Oxford, UK.
Hiby, L. and Krishna, M.B. 2001. Line transect sampling from a curving path. Biometrics
57:727-731.
Karanth, K.U. and Sunquist, M.E. 1992. Population structure, density and biomass of large
herbivores in the tropical forests of Nagarahole, India. Journal of Tropical Ecology
8:21-35.
Kumar, N.S. 2000. Ungulate density and biomass in the tropical semi-arid forest of
Ranthambore, India. M.S. Thesis, Pondicherry University, Salim Ali School of
Ecology and Environmental Sciences, Pondicherry, India.
Thomas, L., Laake, J.L., Strindberg, S., Marques, F.F.C., Borchers, D.L., Buckland, S.T.,
Anderson, D.R., Burnham, K.P., Hedley, S.L. and Pollard, J.H. 2001. DISTANCE
4.0. Beta 3. Research Unit for Wildlife Population Assessment, University of St.
Andrews, Scotland, UK.
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C H A P T E R 11
STATISTICAL CONCEPTS: ESTIMATING ABSOLUTE
DENSITIES OF TIGERS USING CAPTURE-RECAPTURE
SAMPLING
James D. Nichols1 and K. Ullas Karanth2
United States Geological Survey, Patuxent Wildlife Research Center, Laurel, MD-20708-4017, USA.
1
Email: <jim_nichols@usgs.gov>
Wildlife Conservation Society (International Programs), Bronx, New York, NY-10460-1099, USA.
2
Email: <ukaranth@wcs.org>
INTRODUCTION
Capture-recapture can be viewed as an animal survey method in which the count statistic is
the total number of animals caught, and the associated detection probability is the probability
of capture. This probability is estimated using the pattern of captures and recaptures of
animals over the survey period. For example, a typical design might involve a small-mammal
trapping grid. Traps are set and animals trapped on the first sampling occasion are physically
marked (tagged) individually to permit identification in subsequent sampling periods. On the
second sampling occasion (e.g., the next day), animals that represent recaptures (they were
caught at sampling occasion 1) are recorded. New animals are recorded and given individual
marks. The process is repeated for the duration of the study (perhaps five consecutive days).
The data resulting from the study then consist of the total number of animals that were caught
and the record of when (on what sampling occasions) each animal was caught. The total
number of animals caught is the count statistic, and the detection or capture probability is
estimated from the records of the individual capture histories.
Although capture-recapture methods were originally developed for such cases where animals
are physically caught and marked artificially with tags, the underlying concepts are valid
even when animals are identified using natural markings. In the case of tigers (and other
patterned animals such as leopard, chital etc.), we can rely on their distinctive markings in the
form of stripes or spots. Another option would be to identify individuals based on DNA
material obtained from their hair or scat, although this approach is still in its infancy (e.g.,
see Mills et al. 2000). As long as individuals can be identified reliably and sampling methods
provide records of detection and non-detection over the different sampling periods of the
survey, then the resulting data can be used in a capture-recapture framework to estimate
animal abundance.
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 Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling
Recent studies conducted under controlled conditions suggest that a few tigers can be
individually identified from measurements of their tracks, using statistical shape
discrimination software (Riordan 1998). However, the issue is not whether track
measurements obtained on standard substrates and compared among a small
number (<12~17) of tigers yield correct identification of a high proportion of individuals.
Such controlled studies do not address several serious problems associated with field surveys
(Karanth 1999; Karanth et al. in prep.). We are also uncertain how such probabilistic individual
animal identifications can be used in a formal capture-recapture sampling framework for
estimation of abundance.
PHOTO TRAPPING TIGERS
Camera trapping of tigers for the purpose of estimating abundance is described in some detail
by Karanth (1995) and Karanth and Nichols (1998, 2000). The practical aspects of camera
trapping tigers are explained in Chapter 12. The basic methodology involves setting out
camera traps within some area of interest. The configuration of the traps can vary
with the situation, but the main concern is to cover an area fairly completely, in the sense that
it would be difficult for a tiger in the sampled area to travel about and not encounter at least
one camera trap. So the traps should be set in a manner to prevent having “holes” in the
sampled area in which tigers could move without any chance of being “captured”
(photographed).
In an ideal situation, there are sufficient camera traps to cover the entire area of interest with
the available traps. In this case, all traps are set for several consecutive days. Each day is
considered as a sample period. In other cases, the area to be sampled is so large that all
available traps do not cover the area without leaving potential “holes” in coverage. In such a
case, there are various possible schemes for rotating traps to cover the entire area, but in
these cases, the entire area is covered once every two or three days, perhaps. In this sampling
situation, the sampling period is defined by the data collected over the number of days necessary
to cover the entire area. Additional discussion of details associated with sampling designs
will be presented in a subsequent section.
The data obtained from camera trapping are summarized as capture-history data. Each animal
captured at least once during the survey has a capture history. A capture history is simply a
row of 1’s and 0’s indicating on what periods an animal was caught. For example, history
010110 denotes an animal that was caught in periods 2, 4 and 5 and not in periods 1, 3 and 6,
of a 6 - period study.
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Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling
CAPTURE-RECAPTURE MODELS
Capture-recapture models can be viewed as probabilistic expressions describing the process
that gives rise to capture-recapture data. Capture-recapture models are frequently classified
according to requisite assumptions about population “closure”. Closed population models are
used when no gains to, or losses from, the population occur between sampling occasions.
Because of this assumption of no population change, closed models are generally applied to
studies conducted over relatively short time periods. Closed population models permit estimation
of abundance.
Open population models are used when there are gains, losses, or both occurring between
sampling periods. Longer time intervals typically separate sampling occasions.
Such models permit estimation of abundance, and also local survival rate and number of
recruits.
Both approaches can be used with tiger camera trap data. Here our interest is in abundance
estimation, so we focus on closed models (Karanth 1995; Karanth and Nichols 1998). Camera
trap data from long-term studies at Nagarahole, India are currently being analyzed to understand
tiger population dynamics using open models (K.U. Karanth and J.D. Nichols, unpublished
data).
Capture–Recapture Models for Closed Populations
Lincoln-Petersen 2-sample estimator
The Lincoln-Petersen estimator is simple to compute and provides an intuitive basis for
understanding capture-recapture models and estimation (see Seber 1982). In most camera
trapping situations, it would be unwise to trap for only two sample periods (e.g., two days or
nights). However, one could use the two-sample Lincoln-Petersen estimator by aggregating
sample periods into two groups. For example, we could denote days 1-10 as “occasion 1” and
days 11-20 denoted as “occasion 2”.
Define the following statistics obtained directly from the collected camera trapping
data:
n1 = number of animals caught and released (with identity known) on occasion 1,
n2 = number of animals caught and released on occasion 2, and
m = number of animals recaptured in period 2 (these animals were caught at both
occasions 1 and 2).
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 Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling

Define the unknown quantity of interest as: we again obtain the same estimator:

N = total of animals in the sampled population. n + n2 − m n1 + n2 − m nn

Nˆ = 1 = = 1 2 (11.4)
pˆ 1 − (1 − pˆ 1 )(1 − pˆ 2 ) m
Then define the following model parameters:
The final estimator of Equations 11.2-11.4 is known as the Lincoln-Petersen estimator (e.g.,
pi = probability that an animal exposed to sampling efforts in the sampled area is Seber 1982). The following bias-adjusted estimator of Chapman (1951) is recommended for
actual use, especially when sample sizes are small:
captured on occasion i, i = 1, 2, and
p = 1 – (1-p1) (1-p2) = probability that a member of N is caught at least once during ( n + 1)(n2 + 1)
Nˆ = 1 −1 (11.5)
the study. ( m + 1)

An approximately unbiased estimator for the variance associated with the estimator of Equation
Estimation proceeds by first estimating detection probabilities for the two sampling periods.
(11.5) is given by:
This is accomplished by conditioning on the animals captured in one occasion and then asking
what proportion of these were captured in the other occasion as well. For example, in order to ( n1 + 1)(n2 + 1)(n1 − m)(n2 − m)
estimate p1, we condition on the animals caught in period 2 (we know these were present at vaˆr ( Nˆ ) = (11.6)
( m + 1) 2 (m + 2)
period 1, because the population is closed) and ask how many of these were caught in period
1. We estimate p2 in a similar manner and then estimate p using pˆ 1 and pˆ 2 . from Seber (1970, 1982). An approximate 95% confidence interval can be constructed for
the estimate as:
m m
pˆ 1 = , pˆ 2 =
n2 n1 Nˆ ± 1.96 SˆE ( Nˆ )

This approximate confidence interval is based on the fact that Nˆ should be distributed
pˆ = 1 − (1 − pˆ 1 )(1 − pˆ 2 ) (11.1)
approximately as normal with large sample sizes.
C
Recall that the canonical estimator (Equation 3.2) is given as Nˆ = , where C is the count
pˆ As an example of the Lincoln-Petersen estimator, consider a camera trapping study at an
Indian wildlife reserve. Assume that 60 camera traps are placed along trails in tiger habitat
statistic (total number of individual animals caught, in this case), and pˆ is the estimate of the and that photos are taken for two consecutive nights. The following statistics result from this
associated capture probability. If we base estimation on the count statistic from period 1, then study:
we obtain the following estimator:
n1 = 7 = tigers photographed on the first night,
n n1 nn n2 = 5 = tigers photographed on the second night, and
Nˆ = 1 = = 1 2 (11.2)
pˆ 1 m / n2 m
m = 2 = tigers photographed on both nights.
If we instead base estimation on the count statistic from period 2, then we obtain the same These statistics can be used to estimate capture probabilities and abundance.
estimator:
Capture probabilities for each night (sampling occasion) and for both nights combined are
n n nn
Nˆ = 2 = 2 = 1 2 (11.3) estimated as:
pˆ 2 m / n1 m
2
Finally, if we base estimation on the summed count statistic from both periods, 1 and 2, then pˆ 1 = = 0.40
5

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 Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling

2 101, indicating an animal caught at periods 1 and 3 of a 3-period study. Example probability
pˆ 2 = ≈ 0.29 structures for capture history 101 under the above three models are given by:
7

pˆ ≈ 1 − (1 − 0.40)(1 − 0.29) ≈ 0.57 M0 Mt Mb

based on Equation 11.1. The Chapman estimator for abundance is:
(7 + 1)(5 + 1)
Nˆ = − 1 = 15
(2 + 1)
with variance and standard error:
vaˆr( Nˆ ) =
((7 + 1)(5 + 1)(7 − 2)(5 − 2) ) = 20
( 2 + 1) 2 (2 + 2)
SˆE ( Nˆ ) = 20 ≈ 4.47
So the estimated number of tigers in the sampled area is 15. We can compute an approximate
95% confidence interval on this estimate as:
Nˆ ± 1.96SˆE ( Nˆ ) ≈ 15 ± 8.76 = (6.24, 23.76)
Thus, the estimate is not very precise and there is a moderate degree of uncertainty associated
with the estimate. This relative imprecision is to be expected with the small sample sizes that
are likely to characterize studies of rare, secretive animals.
K-sample closed population capture-recapture models
Capture history data from studies with K>2 sample periods permit greater flexibility in
modeling. Models for capture history data are based on parameters associated with the events
that give rise to the capture histories. Models developed for capture histories from K-sample
studies deal with three basic sources of variation in capture probability: Time (sample period),
behavior (trap response) and individual heterogeneity (Otis et al. 1978; White et al. 1982).
These probabilistic models thus incorporate different sets of assumptions about capture
probability.
For example, model M0 assumes that every animal shows the same capture probability (p)
for each sampling period (no variation in p). Model Mt assumes that capture probability
varies by sampling occasion (pi, i = l. .., K). Mb assumes that capture probability differs for
animals that have (c), and have not (p), been caught previously. Consider capture history
p(1-p)p p1(1-p2)p3 p(1-c)c
Model Mh permits each individual in the sampled population to have a different capture
probability.
In addition to these models incorporating single sources of variation in p, there are models
corresponding to all possible combinations of these sources of variation (Mtb, Mth, Mbh, Mtbh).
Estimators for abundance are available under all models, although that for Mtbh requires
ancillary data (on trapping effort). Most estimates are obtained iteratively, so computer
programs are required for estimation. Program CAPTURE (Rexstad and Burnham 1991)
computes estimates under all of these models, and program MARK (White and Burnham
1999) can be used for most models as well.
A closure test is conducted by CAPTURE, and, another test has been developed recently by
Stanley and Burnham (1999). If the closure assumption is rejected, then open models (e.g.,
Pollock et al. 1990) can be used to estimate abundance. The only disadvantage of using open
models is that they do not permit robust estimation of abundance in the face of heterogeneity
or trap response.
Model selection is based on goodness-of-fit and between-model tests. A discriminant function
model selection procedure (based on simulated data) has been built into program CAPTURE.
As a technical note, an information-based model selection criterion known as Akaike’s
Information Criterion, AIC (see Burnham and Anderson 1998) is commonly used for model
selection with other classes of models. However, the closed models that include heterogeneity
cannot be evaluated using standard approaches (different estimation methods must be used
for these), so AIC cannot be used with model sets that include heterogeneity models. This
situation will change soon, as the finite mixture models of Norris and Pollock (1996) and
Pledger (2000) have been incorporated into program MARK (White and Burnham 1999) to
permit use of likelihood methods and computation of AIC for heterogeneity models.
Of all the estimators associated with the different models in CAPTURE, the jackknife estimator
for model Mh tends to be the most robust to deviations from model assumptions. This is
important because the model selection procedure in program CAPTURE requires large sample
sizes in order to perform well. In the future, model weighting will likely be used to develop
estimators (Stanley and Burnham 1998). With model weighting, the final estimate is computed

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 Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling

as the weighted mean of estimates produced by the various models in the model set, with the Density Estimation
weights coming from AIC values or some other measure of the relative support for the different
The above approaches (K-sample models implemented in program CAPTURE, or the 2-
models (e.g., see Burnham and Anderson 1998).
sample Lincoln-Petersen estimator) provide estimates of N and var ( Nˆ ). Sometimes, we are
interested in tiger population density, D = N/A, where A is the area from which animals are
As an example, we consider our camera trapping data from Kanha Reserve in central India
sampled. If entire reserves or other areas with discrete boundaries are sampled, then A may
(Karanth and Nichols 1998). The area was trapped during October – December 1995. There
be known. If portions of parks or fractions of available tiger habitat are sampled, then A will
were 10 sampling occasions totaling 803 trap nights of effort and yielding 59 captures of 26
not be known and must be estimated. One approach to this is to form a polygon by drawing
different individual tigers. Capture histories for this study are presented in Table 11.1.
lines on a map, connecting the locations of all the outermost camera traps. Then add to this
boundary line a “boundary strip” of width corresponding to the estimated distance from
These capture histories were used with program CAPTURE (Rexstad and Burnham 1991) to
which sampled tigers have been drawn. Radio telemetry provides one approach for obtaining
compute statistics related to the closure assumption and to model selection as well as model-
information on distances moved. An ad hoc approach to estimation of boundary strip width
based estimates of abundance. The closure test statistic from program CAPTURE provided
is to use ½ the mean (averaged over all animals) of the maximum distance between capture
no evidence that the closure assumption was violated (z=0.57, P=0.72). Thus, we concluded
locations for the entire study. A simulation study of this and other methods of estimating
that the closed population models of CAPTURE were appropriate for these data and that we
boundary strip width (Wilson and Anderson 1985) indicated that this approach was very
did not need to instead turn to models for open populations such as those of Pollock et al.
reasonable and performed fairly well.
(1990).
Define di as the maximum distance moved between recaptures for animal i, and let m denote
Based on the discriminant function model selection procedure, models M0 and Mh were likely
the number of animals that were caught at least twice during the study. Then the mean and
the most appropriate models for this data set. However, our a priori belief that there would be
variance of maximum distance moved are computed as:
heterogeneity in capture probabilities among individual tigers because of social structure
(Chapter 2) and unequal access to camera traps influenced our choice of the Mh model. Also m

because of the robustness of the jackknife Mh estimator to deviations from model assumptions ∑d i

(Otis et al. 1978), we were more comfortable with the estimates from this model. Estimates dˆ = i =1

m
under the above two models are presented below.
m

Capture probability estimate Abundance ∑ (d i − dˆ ) 2

vaˆr(dˆ ) = i =1 (11.7)
Model per occasion entire study Nˆ SˆE ( Nˆ ) 95%CI m( m − 1)

M0 0.21 0.93 28 2.04 26-33 Boundary strip width, W, and its variance, can be estimated as:
Mh 0.18 0.79 33 4.69 29-49
dˆ
Wˆ =
2
The estimates under these two models differ in predictable ways. First, in the presence of
heterogeneity, estimates under model M0 should be negatively biased and smaller than estimates
vaˆr( dˆ )
under Mh (Otis et al. 1978; Pollock et al. 1990). This expectation is reflected in the above vaˆr(Wˆ ) = (11.8)
estimates under the two models. Second, estimates under model Mh are expected to be much 4
less precise than those under model M0, and this expectation is reflected in the above estimates The area of the polygon defined by the outermost traps plus the estimated boundary strip can
of standard error. be obtained manually using a map and a planimeter or using GIS (see Karanth and Nichols

128 129
 Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling

1998). However, note that this trap polygon is not the same as the Minimum Convex Polygon The final quantity required for Equation 11.9 is then vaˆr( Aˆ (W )) , which can be estimated in
often used in radio telemetry studies. Denote this estimated area (often referred to as effective either of two ways. One approach is based on the bootstrap (Efron and Tibshirani 1986) and
area) as Aˆ (W), indicating the area obtained by adding to the area covered by traps, involves selecting m ' maximum distances moved with replacement from the m actual distances.
the boundary strip of estimated width Wˆ (see Figure 11.1). Then estimate density and its These distances are used in conjunction with Equations 11.7 and 11.8 to obtain an estimate of
variance as: Wˆ * , where the “*” superscript denotes that this is an estimate from a bootstrap replicate.
This Wˆ * is then used to create a new area, Aˆ (W * ) , which is determined by planimeter or GIS
Nˆ methodology. A replication-based variance estimate is then computed from these bootstrap
Dˆ =
Aˆ (W ) replicates, Aˆ (W * ) , and used as an estimate of var ( Aˆ (W ) ).

The other approach is to approximate the shape of the sampled area with a circle of radius
 vaˆr( Aˆ (W )) vaˆr( Nˆ ) 
vaˆr( Dˆ ) = Dˆ 2
+  (11.9) c + W, where c is a constant computed to satisfy the equality:
ˆ 2
Nˆ 2 
 [ A(W )]
Aˆ (W ) = π (c + Wˆ )
2

where vaˆr( Nˆ ) is the model-based sampling variance and is computed for the selected estimator
by program CAPTURE.
A delta method (see Seber 1982) approximation of vaˆr( Aˆ (W )) can then be computed as:

vaˆr( Aˆ (W )) = 4π (c + Wˆ ) vaˆr(Wˆ )
2 2

This approach to density estimation was described in detail by Karanth and Nichols (1998).

As an example, consider the camera trapping data from Kanha Reserve used in the previous
example. The average maximum distance moved for animals captured at least twice was
dˆ = 4.31 km, yielding an approximate strip width of:

dˆ 4.31
Wˆ = = ≈ 2.16 km.
2 2
The area of the polygon obtained by connecting the outermost camera traps was 142.37 km2.
The addition of the Wˆ =2.16 approximate strip width to this polygon yielded an effective area
of Aˆ (W ) = 282.02 km2. The density is then estimated by dividing the estimated population
size ( Nˆ = 33 ; see previous example) by the effective area:

Nˆ 33
Dˆ = = ≈ 0.117
Aˆ (W ) 282.02

which is a density of 11.7 tigers per 100 km2. Although we do not detail the computations for
the standard error here, the resulting estimate is SˆE ( Dˆ ) ≈ 0.0193 or ±1.93 tigers per 100
Figure 11.1 - A map of Kanha Reserve, India, illustrating the layout of camera trap points and the km2.
effective sampled area (Source: Karanth and Nichols 2000).

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 Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling
SURVEY DESIGN CONSIDERATIONS
An overview of the sampling associated with camera trapping was provided above. Here we
provide general considerations about temporal and spatial aspects of sampling.
Time – A general objective in camera trapping is to keep sampling time “short” relative to
tiger population turnover so that closed models can be used. However, the definition of “short”
will be situation-dependent and will likely be determined by the logistics of each sampling
situation. It would be excellent, but likely impossible, to complete sampling within a 2-week
period. Periods of 4-6 weeks will more likely be required, and 8-12 weeks could perhaps be
used. Longer time intervals increase the likelihood of models for open populations being
needed. Closed models are preferred for reasons of precision and estimator robustness, but if
closed models cannot be used, then open models are available (Seber 1982; Pollock et al.
1990).
Space – The primary objective to keep in mind when considering space is that at each capture
“occasion”, every animal in the study area should have some non-negligible chance of being
captured. Even model Mh does not deal well with animals characterized by capture probabilities
that approach 0. It is desirable to avoid the situation where a tiger within the study area can
have its entire range lie within an area that does not overlap at least one or two camera traps.
Capture probabilities should also be relatively similar among the animals in the sampled area,
to the degree possible.
Possible approaches to dealing with space and time – Below, we present four
possible approaches to dealing with these space-time considerations in camera trapping
studies.
(1) If an adequate number of camera traps is available and if the study area is sufficiently
small, then traps can be spread throughout the area once and then checked each day for
say 5-30 consecutive days. This approach yields standard capture history data.
(2) Frequently there will not be enough traps to sample the entire area of interest, but very
frequent (e.g., daily) movement of traps to any potential trap site within the area may be
logistically possible. In such a situation, we recommend dividing the area of interest into
a grid system. We emphasize here that the “grid” system for photo trapping tigers need
not be a typical square grid pattern that is used in capture-recapture studies of rodents,
but is simply a system of subdividing the study area into a number of potential sampling
units, where the size of a unit is defined as an area so small that you would not want two
traps placed there at the same time. On day 1, randomly select y “grid cells”, where y
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Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling
corresponds to the number of available camera traps. Place traps at promising sites within
each cell. On day 2, randomly select y new cells, pick up cameras and move them to
promising trap sites within the new cells and set them. Repeat this procedure for 5-30
days. This approach again yields standard capture history data.
(3) In other situations, not only will the number of traps may not be adequate to sample the
area of interest, but also the traps cannot be readily moved to any location within the area
of interest. However, moderately frequent movement of traps will be logistically possible,
especially if movement is to nearby locations. Assume that ¼ of the area can be covered
with traps at any one time. In this case, we can divide the study area into four trapping
blocks. The investigator might place traps in block 1 for 1-5 days, then in block 2 for
1-5 days, next in block 3 for 1-5 days and finally in block 4 for 1-5 days. This entire
set of days will be denoted as sample occasion 1. Then the procedure is repeated, and the
resulting captures/recaptures will be assigned to capture occasion 2. The cycle is repeated
until a minimum of 5 “occasions” are obtained. Capture data are combined over all
blocks for days 1-5 to define occasion 1 (e.g., the first “cycle” defines occasion 1). Captures
and recaptures are combined over all 4 blocks for the second occasion (e.g., the
second cycle defines occasion 2), etc. The resulting capture histories are then used
in the analyses.
(4) Finally, we can envision situations where the number of traps is inadequate to sample the
area of interest and frequent movement of traps is not logistically possible. Again assume
a situation where ¼ of the area can be covered with available traps. The traps can be set
out in the first block and left for 5-20 days. Traps are then moved to block 2 and left for
the same number of days as block 1. The procedure is repeated for blocks 3 and 4. The
number of captures for capture occasion 1 is obtained as the total number of captures
occurring on the first day of trapping in each block. The number of captures/recaptures
for capture occasion 2 is obtained as the sum of captures/recaptures for the second capture
day at each block, and so on. This approach again yields capture history data that can be
used with program CAPTURE.
Approaches 1 and 2 are clearly the most desirable and approaches 3 and 4 less desirable from
the perspectives of modeling and estimation. For example, time variation will be difficult to
deal with under approaches 3 and 4, because each capture occasion covers multiple calendar
days. Thus, it no longer makes sense to think of each occasion as characterized by a certain
set of environmental conditions for a particular day. So the “M t” models are not as likely to be
useful under sampling design approaches 3 and 4. Nevertheless, all of the above approaches
should “work” for tigers and similar species.
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 Statistical Concepts: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling
CONCLUSION
Camera trapping seems to provide a reasonable and rigorous way to estimate relative and
absolute abundance of tigers. It clearly works better in high-density situations than in places
where tigers are sparse. It is logistically most easily employed in areas with good road and
trail systems but can be used in remote situations as well (Chapter 12). Finally, we note that
camera trapping repeated over time (e.g., once per year) can provide a rigorous monitoring
program and permits use of Pollock’s (1982) robust design to estimate survival rate, rate of
population change, year to year population turnover, etc. (K.U. Karanth and J.D. Nichols
unpublished data).
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Otis, D.L., Burnham, K.P., White, G.C. and Anderson, D.R. 1978. Statistical inference from
capture data on closed animal populations. Wildlife Monographs 62:1-135.
Pledger, S. 2000. Unified maximum likelihood estimates for closed capture-recapture models
for mixtures. Biometrics 56:434-442.
Pollock, K.H. 1982. A capture-recapture design robust to unequal probability of capture.
Journal of Wildlife Management 46:757-760.
Pollock, K.H., Nichols, J.D., Brownie, C. and Hines, J.E. 1990. Statistical inference for
capture-recapture experiments. Wildlife Monographs 107:1-197.
Rexstad, E. and Burnham, K.P. 1991. User’s guide for interactive program CAPTURE.
Abundance estimation of closed animal populations. Colorado State University,
Fort Collins, CO, USA.
Riordan, P. 1998. Unsupervised recognition of individual tigers and snow leopards from their
footprints. Animal Conservation 1:253-262.
Seber, G.A.F. 1970. The effects of trap response on tag-recapture estimates. Biometrika
26:13-22.
Seber, G.A.F. 1982. The estimation of animal abundance and related parameters. MacMillan,
NewYork, NY, USA.
Stanley, T.R. and Burnham, K.P. 1998. Estimator selection for closed-population capture-
recapture. Journal of Agriculture, Biology and Environmental Statistics 3:131-150.
Stanley, T.R. and Burnham, K.P. 1999. A closure test for time-specific capture-recapture
data. Environmental and Ecological Statistics 6:197-209.
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White, G.C., Anderson, D.R., Burnham, K.P. and Otis, D.L. 1982. Capture-recapture removal Table: 11.1. Capture histories of individual tigers from camera trapping at Kanha Reserve,
methods for sampling closed populations. Los Alamos National Laboratory India during October-December 1995.
Publication LA-8787-NERP. Los Alamos, NM, USA.
Tiger Sampling occasion
White, G.C. and Burnham, K.P. 1999. Program MARK: Survival rate estimation from both identification 1 2 3 4 5 6 7 8 9 10
live and dead encounters. Bird Study 46:120-139.
number
Wilson, K.R. and Anderson, D.R. 1985. Evaluation of two density estimators of small mammal KNT–101 1 0 0 1 0 0 0 1 1 0
population size. Journal of Mammalogy 66:13-21. KNT–102 1 0 0 0 0 0 0 1 0 1
KNT–103 1 1 0 1 0 0 0 0 1 1
KNT–104 0 1 1 0 0 0 0 1 1 1
KNT–105 0 1 0 1 0 0 0 0 0 1
KNT–106 0 1 0 0 0 0 0 0 0 0
KNT–107 0 0 1 0 0 1 1 0 0 0
KNT–108 0 0 1 1 0 0 0 0 0 0
KNT–109 0 0 0 1 0 0 0 0 1 1
KNT–110 0 0 0 1 1 0 0 1 1 0
KNT–111 0 0 0 1 0 0 1 0 0 1
KNT–112 0 0 0 0 1 0 0 0 0 0
KNT–113 0 0 0 1 0 0 0 0 0 0
KNT–114 1 0 0 1 0 0 0 0 0 0
KNT–115 0 0 0 1 0 0 0 0 0 0
KNT–116 0 0 0 1 1 0 0 0 0 0
KNT–117 0 0 0 1 0 0 1 0 0 0
KNT–118 0 0 0 0 1 1 0 0 0 0
KNT–119 0 0 0 0 1 0 0 0 0 0
KNT–120 0 0 0 0 0 1 0 0 0 0
KNT–121 1 0 0 0 1 0 1 0 0 0
KNT–123 0 1 0 0 0 0 0 0 0 0
KNT–124 0 0 0 1 0 0 0 0 0 1
KNT–125 0 0 0 0 0 1 0 0 0 0
KNT–126 0 0 0 0 0 0 0 0 0 1
KNT–127 0 0 0 0 1 0 0 0 0 0

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 C H A P T E R 12

FIELD SURVEYS: ESTIMATING ABSOLUTE DENSITIES OF

TIGERS USING CAPTURE-RECAPTURE SAMPLING

K. Ullas Karanth1, N. Samba Kumar2 and James D. Nichols3

Wildlife Conservation Society (International Programs), Bronx, New York, NY-10460-1099, USA.
1

Email: <ukaranth@wcs.org>
2
Centre for Wildlife Studies, 26-2 Aga Abbas Ali road (Apt. 403), Bangalore, Karnataka-560 042, India.
Email: <wcsind@bgl.vsnl.net.in>

United States Geological Survey, Patuxent Wildlife Research Center, Laurel, MD-20708-4017, USA.
3

Email: <jim_nichols@usgs.gov>

INTRODUCTION

BLANK In Chapter 11, we saw that sampling animal populations by repeatedly ‘catching’ identified
individuals can generate capture histories. From these histories, capture frequency statistics
and estimates of capture probabilities can be derived. Estimates of capture probabilities permit
us to estimate the abundance and density of animals in the surveyed area, without catching all
individuals in the population.

In traditional capture-recapture studies (mark-recapture), target species such as rodents, birds

and fish are usually marked using artificial tags or bands. However, it is not easy for us to
catch and mark tigers in this manner! Fortunately, tigers have natural body markings in the
form of stripes, which allow us to identify individuals. If we can ‘catch’ individually identifiable
tigers repeatedly in samples drawn from a wild population, we can then build capture histories,
estimate capture probabilities and thereafter estimate tiger numbers and densities using powerful
capture-recapture models. At this point in time, there are two such ways of non-invasively
‘catching’ individual tigers from a population: ‘Photographic captures’ and ‘DNA-based
captures’.

In photo trapping methods, the tigers themselves trigger self-activated camera traps, and
take their own pictures. In the DNA-based sample surveys, instead of a picture of the
animal, the individual identification is provided by a sample of body hair that is snagged
by a device placed in the tiger’s path. DNA material extracted from the tissue at the root
of the hair is then used to identify individuals using advanced laboratory techniques.
Techniques under development may soon make it possible to extract DNA material from tiger
scats also.

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At the present time, DNA-based methods for identifying individual tigers are not fully
developed, although they have been used to estimate numbers of grizzly bears and
other species using capture-recapture protocols (Woods et al. 1999; Mowat and
Strobek 2000). Therefore, in this chapter we will write only about the capture-recapture
sampling of tigers using camera trap photos. However, many of these ideas will be applicable
to DNA-based field surveys if and when they become cost-effective and practical
survey tools.

CHOICE OF CAMERA TRAP EQUIPMENT

The camera trap equipment for photographing tigers has two components: The camera and
a tripping device that fires it. Although professional photographers sometimes use
expensive single-lens-reflex, digital or video cameras to get high quality pictures, many ordinary
electronic cameras can be hooked up to tripping devices to get usable pictures of tigers.
However, such cameras must have auto-focus and auto-rewind capabilities and internal
circuitry that permit shooting several pictures in succession using flash without exhausting
the battery. The camera should withstand exposure to moisture, extreme temperatures and
rough field use, and its batteries should last for several days, even with the shutter cover
kept open.
To conserve the battery power, most modern electronic cameras go to ‘sleep’ if the shutter
cover is left open for a couple of hours or so. With such ‘idiot-proof’ cameras, the tripping
device should be able to periodically ‘wake the cameras up’ electronically. Otherwise, the
picture of the first animal that ‘wakes up’ a sleeping camera will be missed.
Several different types of tripping devices can be used in camera traps. In some units two
metallic strips held apart inside a pressure pad come into contact with each other when the
tiger steps on the pressure pad, completing an electrical circuit that fires the camera.
More commonly, electronic tripping that employs an infrared device is used. There are two
kinds of such infrared tripping devices. The ‘active’ type units have an infrared transmitter
that emits a beam that is received by an infrared receiver placed opposite to it. When the tiger
walks between the two units, the beam is interrupted for a few seconds, completing a circuit
that fires the camera (Figure 12.1). On the other hand, ‘passive’ infrared type devices ‘sense’
the heat emanating from the body of the tiger passing in front of it. The sensor then completes
a circuit that fires the camera. Among the well-known commercial brands of camera traps,
Trailmaster 1500 units use active infrared tripping, whereas Trailmaster 1000 and CamTrakker
units use passive devices.
Figure 12.1 - An active infrared camera trap unit, showing the main components and set up.
Although camera traps can be homemade devices that work in a variety of ways, commercially
made units tend to be more reliable. If your goal is to conduct a proper survey with
effective coverage of a large area, requiring many units deployed under a rigorous
sampling schedule (Chapter 11), it is better to invest in good commercially made camera
traps. Appendix 6.0 lists the names of some manufacturers. Additionally, the following
factors must be considered when choosing equipment for a field survey aiming to estimate
tiger density:
1. You need unambiguous identification of individual tigers to build their capture histories.
The stripe patterns on the left and right flanks of the same tiger differ. Therefore, to
identify any particular individual with certainty, both of its sides must be photographed
simultaneously, at least once during the survey to link the two profiles. In practical terms
this means that every camera trap unit must have two cameras positioned opposite to each
other. Because most pictures are taken at night, a built in electronic time delay (of 30-40
milliseconds) that prevents the two flashes from flaring the pictures is useful.
2. In some makes, a single tripping device fires two cameras through a multi-camera trigger.
In others, two separate tripping units need to be bought to operate the two cameras. The
relative costs need to be considered keeping this factor in view.

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3. Generally, passive infrared units tend to be less prone to false tripping from moisture
condensation, insects, and vibration etc., compared to active type infrared units. In areas
where rain is a major problem, this is a very important advantage of passive infrared units.
4. It is generally easier to compose the picture and fix the tiger’s anticipated position with
active infrared units, because of the relatively narrow electronic beams that they generate.
5. The minimum time delay that can be set between two successive pictures tends to be much
shorter (a few seconds) in active units compared to passive units (several minutes). The
practical implication of this difference is that, if two or more tigers are traveling together,
only the first animal is likely to be photographed by the passive unit, whereas the active
unit is likely to get others also.
underfoot conditions, sandy streambeds, and forest roads make good ‘trap lines’ along which
camera traps can be set.
Once a trap site is chosen, the goal is to frame the picture so that the tiger’s flanks are clearly
photographed. Although stripes on any part of the tiger’s body can be used for identification,
broadside pictures are the easiest to use.
The path should not be modified drastically and the traps should be placed unobtrusively and
cryptically. Often a simple trick like throwing a few branches around a trap can prevent it
from sticking out like sore thumb. Masking tape of a dull color should be used on metallic or
brightly colored parts. All loose cables should be hidden under leaf litter or soil.

6. If cameras that go into ‘sleep’ mode are used as mentioned earlier, the tripping device must
include the circuitry that ‘wakes up’ the cameras periodically to prevent potential loss of
pictures of tigers.
7. Some manufacturers enclose their units in moisture-resistant containers. In rainy or humid
areas this is an important advantage. Sometimes, other animals like bears and elephants
damage the units or humans may steal or vandalize them. In such situations, a rugged
protective shell may need to be deployed to protect the cameras. Appendix 5.0 shows a
protective shell designed by J. Amarnath that we have found useful.
The two cameras should be positioned about 7 meters apart, on either side of the tiger’s
anticipated path. Even if the animal walks on either edge of the path, usable pictures are still
obtained at this distance. In active infrared units, the electronic beam should be set 45 cm
above the ground to catch both adults and cubs (Figures 12.2 and 12.3). In Trailmaster traps,
we found that setting the period of beam interruption to 5 missing pulses is useful for catching
tigers while avoiding film wastage on small animals.

CONDUCTING CAMERA TRAP SURVEYS

Choice of Sites and Setting up Traps

Given that tigers occur at low densities of 10-20 animals/100 km2 even at the best sites,
getting a ‘photographic capture’ of a tiger is a rare, uncertain, event. Therefore, more than
anything else, the camera trap sites must be chosen to maximize capture probabilities: They
should not be randomly selected spots in the sampled area (see Chapter 11). Tigers regularly
use travel routes (Chapter 2) and do not move randomly through space. If traps are set out
randomly, most sites are likely to have zero or near zero probabilities of catching a tiger. True
random selection of trapping points will typically yield no tiger captures!

Choice of a good trap site depends on careful preliminary reconnaissance surveys to examine
the presence of tiger and prey signs (tracks, scrapes, scent, scats) indicating past use. More
generally, it depends on your ability to ‘think like a tiger’: Prior knowledge of local naturalists
and hunters is very useful in this task. Convergence of game trails, presence of salt-licks and
water that attract prey animals, often indicate a good site. Usually game trails with soft Figure 12.2 - Setting up a camera trap unit in the field.

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 Field Surveys: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling
Figure 12.3 - A Tiger tripping the camera trap.
Trap Spacing and the Number of Traps
In any capture-recapture sampling scheme (Chapter 11), the spacing of traps is dictated by
the biology of the species. To trap mice the spacing would be scaled to a few meters, for deer
we may use a spacing of a hundred meters or so, and for landscape animals like tigers the trap
spacing has to be set in terms of kilometers. Recall that our goal is not to have ‘holes’ in
which an animal present in the surveyed area has zero or very low probability of being
captured.
In absolute terms what should be the spacing between traps? The distance between traps
must be relatively small, if the minimum tiger home range size is expected to be small. If the
expected home ranges are large, traps can be set farther apart. From previous studies of tiger
social organization (Chapter 2) we know that among all the age sex classes in a population,
adult tigresses that breed have the smallest home ranges (Chapter 2). Therefore, if we ensure
that at least 2-3 traps are placed within an area of the size of the expected female home range
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Field Surveys: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling
(Otis et al. 1978, Karanth and Nichols 1998), individuals in other age-sex classes will
automatically be exposed to greater numbers of traps.
Home ranges of adult tigresses in prey-rich areas can be as small as 10-15 km2, but not any
smaller (Sunquist 1981; Smith 1993; Sunquist et al. 1999; Karanth and Sunquist 2000). In
such areas, camera traps can be set about 2-3 kilometer apart from each other, ensuring that
an adult tigress is potentially exposed to at least 2-3 traps (Figure 12.4). At the other extreme,
in areas where the prey base is poor (temperate zone forests, primary rainforests, over-hunted
sites), the female ranges can be as large as 200-500 km2 (Miquelle et al. 1999). In such areas
camera traps can be deployed 5-10 kilometers apart. We suggest the above figures only as
broad guidelines. Specific local knowledge about tiger movement patterns and home range
sizes are critical considerations while deciding on the trap locations for any survey.
If more than 2-3 traps are placed within an animal’s home range, then it is more likely that the
animal will be caught in each sampling effort. So, within an area of the expected home range
size, it is better to place ‘more traps’ than ‘fewer traps’. However, on the other hand, our goal
also is to catch as many different individual tigers as possible within the area of interest. The
sample size in a capture-recapture survey is the number of individuals caught (Otis et al.
1978; White et al. 1982). Given a fixed number of camera traps at our disposal, we can
potentially increase the number of individual tigers we can catch by increasing the trap spacing,
thereby sampling a larger area that holds more tigers. Therefore, trap spacing becomes a
compromise between these two needs.
Ideally the study area should be reconnoitered beforehand to locate and map several suitable
trapping sites. Thereafter, you can experiment on a map, using different choices of trap sites,
to optimize trap spacing and locations. After several potential trap sites are identified and
mapped, you can choose one of the sampling schemes from among the different designs
explained in Chapter 11. In this process, logistical factors such as the number of camera
traps available, manpower, terrain, mode of transport, all of which determine how often traps
can be set up, checked and moved around become important. While deciding on trap locations,
factors such as weather, animal damage, vandalism and theft, also have to be considered.
Sampling Periods and Time Frame for Camera Trapping
Any camera trap sample survey scheme must include two or more samples (sampling
occasions). These discrete sampling occasions (secondary sampling periods), add up to form
the overall trapping session (primary sampling period). The length of the primary sampling
period is guided by both theoretical and practical considerations. Recall that the critical
parameter we are trying to estimate is the number of tigers present in the surveyed area
(absolute abundance). This tiger population should be ‘geographically closed’ and
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‘demographically closed’ (see Chapter 11) throughout the primary period. Therefore, ideally
the survey should last only a few days, as the closure assumption is more likely to be met for
such short periods.

Generally, if there are more secondary sampling periods, we can use more refined models
and estimators for data analysis (Otis et al. 1978). Having 5-15 sampling periods is better
than having just 2. Even if we have enough traps on hand to cover the entire area in just one
day, 15 days of sampling will be required to attain 15 secondary sampling periods. Usually,
camera traps are not available in such large numbers and one secondary sampling period - or
each ‘pass’ over the entire surveyed area (one pass covering all trapping points once) - takes

Figure 12.4 - Camera trap spacing patterns for sampling a tiger population.
more than one day, depending on the area covered and the sampling design used (Chapter 11).

For example, Karanth and Nichols (2000) used about 60 trap locations at several study
areas of 150-300 km2. They had 15-20 camera trap units on hand at any given time,
and therefore needed 3-4 days to complete each secondary sampling occasion. Therefore,
their primary sampling period ranged from 45 to 60 days to obtain 15 sampling periods.
Instead, if they had decided to obtain only 10 samples, their surveys could have been
completed in 30-40 days. Because tigers are relatively long-lived animals (compared to rodents),
assuming demographic closure for 45-60 days of camera trapping may not be unrealistic.

We note that fixing the duration of the primary sampling period involves a trade-off between
increasing the number of secondary samples (which is good) and the prospect of the closure
assumption being violated (which is bad).

However, camera trapping an area continuously over months and years - as some investigators
appear to be doing - certainly violates the closure assumption. It is incorrect to apply closed
capture-recapture models to the data from these open populations. Although open models can
be used to estimate abundance in such cases, these estimators are not as robust as those based
on closed models. This is particularly true for tigers whose social organization pattern (Chapter
2) almost certainly results in heterogeneity of capture probabilities among individuals. While
closed capture-recapture models can deal with such heterogeneity, open models cannot.
Therefore, open models will yield estimates of tiger abundance that are certainly better than
simple counts of total individuals caught, but are less reliable than estimates based on closed
models.

Data Collection Forms and Protocols

Meticulously following field protocols is important to prevent all sorts of inaccuracies
creeping into the identifications of individual tigers and their capture-histories. We make the
following recommendations:

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 Field Surveys: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling
1. Camera traps have to be checked on a daily basis (or as frequently as possible), to verify
proper working and to record any tripping that occurred previously.
2. A standard data form (specimen shown in Appendix 4.0) rather than field notebooks
should be used to record trapping data. Unique identification numbers should be given to
each individual camera and each tripping unit for easy troubleshooting when these pieces
of equipment malfunction.
3. Individual tigers must be identified correctly from photographs of both flanks. Even if you
decide to use single frontal photos instead, half the data will be lost when the tigers travel
in the opposite direction. Investigators using single camera trapping units will inevitably
lose roughly 50% of the data. Given all the other major expenses and effort involved in
camera trapping tigers, this is inefficient and makes little sense.
4. It is important to uniquely number each film roll before it is loaded. Such identification is
critical for accurately fixing the location and the sampling period for any capture event,
and for matching left and right profiles of tigers for correct individual identification.
Because commercial labs mix up several rolls of film during processing, you must ensure
that each roll you submit is processed under a unique identifying label. If identities of rolls
are mixed up during processing, capture history data you construct can be seriously flawed.
It will also be impossible to match left and right profiles of tigers. If the processed negative
film is reversed while printing also, the tiger identifications will get vitiated. Therefore,
each film roll has to be individually ‘tracked’ from the time it is loaded into a camera until
it is processed and printed.
5. When you check camera traps, pay particular attention to: Battery status; number of
remaining exposures; alignment of the infrared beam and proper mounting of the camera.
You also need to make sure that the date, time, photographing time zone, camera delay
and the number of pulses missed, are all set correctly, depending on the make of the unit.
The date and time of each exposure as read by the camera’s data-back should be carefully
double-checked with those recorded by the tripping device.
6. All the camera trap equipment should be kept free of dust and moisture and maintained as
recommended by the manufacturer’s manual. Fresh battery cells should be used in the
cameras and tripping devices to ensure that data are not lost.
Identifying Tigers from Camera Trap Photos
After the film is processed, the first step is to link the left and right profiles of individual
tigers relying on the data in the field form (Appendix 4.0). The next step is to identify each
individual tiger by comparing two photographs of the same flank.
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Field Surveys: Estimating Absolute Densities of Tigers Using Capture-Recapture Sampling
First, look for key diagnostic features in the stripe pattern like a peculiarly shaped stripe or
other unique features among the large stripes on the flanks. Compare the shape of specific
individual stripes and their positions on the animal’s body. If necessary, use additional stripe
patterns on the limbs, tail and face as supplemental evidence (Figure 12.5).
Figure 12.5 - Identifying individual tigers based on stripe patterns.
After all the individual tigers are identified and given unique numbers, every capture of each
individual tiger is assigned to a particular secondary sampling occasion (Chapter 11) based
on the date and time of capture. We strongly recommend maintaining a spreadsheet database
of all capture records. From this database, the capture histories of each individual tiger can
be placed in capture history matrix format quite easily. In this format, each animal detected at
least once represents a row in the matrix, with each column representing a sampling occasion
and containing either a 1 (indicating capture on that sampling occasion) or a 0 (indicating no
capture). Table 11.1 shows an example of a typical capture history matrix constructed from
camera trapping in Kanha Reserve, India in 1995.
ANALYSIS OF CAPTURE-RECAPTURE DATA
The capture history data described above can be analyzed using computer programs such as
CAPTURE (Rexstad and Burnham 1991) or MARK (White and Burnham 1999) for estimating
the number of tigers within the sampled area. The final analysis of closed model capture-
recapture survey data using program CAPTURE consists of the following steps:
1. Checking results of the closure test to see if the assumption of population closure during
the survey was violated.
2. Checking the results of the various between-model statistical tests, and, the overall model
selection statistic to decide the appropriate model for the data that you have collected.
Often, the statistical tests indicate that the null model M0 (which assumes equal capture
probabilities for all tigers, no trap response, and, no time effects on trapping) is the best-
fit model for your data. However, because we know that tigers are territorial animals
(Chapter 2) and that the estimator under model M0 is not robust to heterogeneity of capture
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tiger and its conservation needs. Pages 5-18 in Riding the tiger: Tiger conservation K. Ullas Karanth1 and James D. Nichols2
in human-dominated landscapes (Eds: J. Seidensticker, S. Christie and P. Jackson).
Cambridge University Press, Cambridge, UK. Wildlife Conservation Society (International Programs), Bronx, New York, NY-10460-1099, USA.
1

Email: <ukaranth@wcs.org>

White, G.C., Anderson, D.R., Burnham, K.P. and Otis, D.L. 1982. Capture-recapture and United States Geological Survey, Patuxent Wildlife Research Center, Laurel, MD-20708-4017, USA.
2

Email: <jim_nichols@usgs.gov>
removal methods for sampling closed populations. Los Alamos National Laboratory
Publication LA-8787-NERP. Los Alamos, NM, USA.
THE NATURAL ADVANTAGE
White, G.C. and Burnham, K.P. 1999. Program MARK: Survival rate estimation from both
live and dead encounters. Bird Study 46:120-139.
Tigers are naturally marked animals. This biological fact confers a singular advantage when
you try to monitor their populations. The purpose of this chapter is to provide a summary of
Wilson, K.R., and Anderson, D.R. 1985. Evaluation of two density estimators of small mammal
how best you can maximize this advantage. Most of the points we make here are relevant to
population size. Journal of Mammalogy 66:13-21.
the studies of other species of naturally marked animals as well. Below are presented some
questions field biologists frequently ask (or should be asking) before they start a program for
Woods, J.G., Paetkau, D., Lewis, D., McLellan, B.N., Proctor, M. and Strobeck, C. 1999.
monitoring tiger populations. However, please note that these brief questions and answers are
Genetic tagging of free-ranging black and brown bears. Wildlife Society Bulletin
not a substitute for reading the rest of this manual!
27:616-627.
How should I begin to think about doing a tiger survey?

Don’t reinvent the wheel: Reading the relevant literature and thinking clearly about your
goals is the best way to start. Nichols (1992) provides an overview of animal population
sampling methods that rely on marked animals. This manual builds on the basic ideas of
marked animal studies and directs you to more advanced resources that are already available.

Why am I doing this tiger survey?

Although they may not formulate these goals clearly in their own minds, biologists, managers
and conservationists monitor tigers specifically to meet one or more of the following four
objectives.

1. To estimate the spatial distribution of tigers (presence or absence) with one of the following
two goals: (A) to determine and map all the locations where wild tigers are present;
(B) to estimate the proportion of habitat occupied by wild tigers or their breeding
populations.

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 Monitoring Tiger Populations: Why Use Capture-Recapture Sampling?
2. To compare the relative abundance of tigers at two different points in time or space: Are
there more tigers in park A or in park B? Or, are there more or fewer tigers in reserve X
now compared to five years ago?
3. To estimate the absolute abundance of tigers (number of tigers in a reserve or population
size) or its derivative, the tiger density (the number of tigers/100 km 2).
4 To estimate annual survival and recruitment rates in a tiger population over the years:
How many tigers in a population survived each year, and how many new tigers were
added to that population?
At the very beginning it is important to decide what your specific objectives are among the
above four, and assess your ability to achieve these in relation to the resources available to
you (Chapter 1). Note that the above objectives are best developed in conjunction with the
overall scientific and/or management questions of interest.
Okay, let us say my goal is to go “everywhere” to simply determine if tigers are present or
not? (Objective 1A above).
Good luck. Usually no survey method can tell you with 100% certainty that tigers are absent
in an area even if you did not find any evidence of their presence. However, you can estimate
the probabilities of not finding sign, given that animals are present, using the general sampling-
based framework described in Chapters 3 and 4. These estimates then translate into estimates
of the proportion of area occupied by tigers.
Why should I consider the sampling-based, model-driven, formal framework you advocate
in this manual for my tiger survey, rather than my own ad hoc method, if my objectives are
1B, 2 or 3?
The simple fact is that you cannot be everywhere at the same time, nor can you be sure of
counting all individual animals in your study areas (Chapter 3). Therefore, whatever you
think you are doing, you end up getting a ‘sample count’ of some sort. The better (less biased
and more precise) your sampling methods are, the more reliable your results will be (Chapters
3, 5, 7, 9 and 11). The approach we recommend allows you to get the most out of your field
data. This approach yields not only estimates of the quantities of interest but also measures of
precision that permit us to judge reliability of our estimates.
When should I use camera traps in my field surveys for tigers? When should I use surveys
of tiger sign like scats or tracks instead of deploying camera traps?
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Monitoring Tiger Populations: Why Use Capture-Recapture Sampling?
If your objectives are 1 and 2, use camera traps only when tiger sign (tracks, scats) surveys
are even more difficult to conduct (in terms of labor, time, money etc.) than camera trapping,
or, in the case of objective 2, if you are especially concerned about bias in estimates of trend
or relative abundance. Use camera traps when your goal is to estimate absolute abundances
or derive less-biased estimates of relative abundances of tigers (objectives 2 and 3) and/or to
estimate survival and recruitment rates (objective 4).
Often you may get more ‘bang for the buck’ by counting tiger signs rather than by using
camera traps. Precision of an index to tiger density is largely a function of sample size:
Encounter surveys for tiger sign may sometimes yield larger samples (e.g., number of tiger
track sets seen/100 km walked) compared to camera trap surveys (number of individual
tigers photographed/100 trap nights).
The local statistician does not recommend the capture-recapture framework you recommend
here: Instead he/she asks me to follow X, Y, Z protocols in my tiger survey.
Estimation of animal population parameters is a very specialized branch of biostatistics.
Often, the approaches of conventional statistics are not highly relevant to animal population
surveys. If the local statistician is willing to become closely involved with the study, then that
person may be able to develop useful estimation approaches. However, if the local statistician
provides standard consulting advice and does not happen to be an expert in wildlife statistics,
then we recommend our approach as more likely to be useful. Please refer to the advanced
literature for details (Thompson 1992; Buckland et al. 1993; Lancia et al. 1994; Thompson
et al. 1998; Johnson 1999; Yoccoz et al. 2001; Williams et al. 2002).
Significant advances have occurred in the approaches to animal population estimation during
the last several decades. Not all the statisticians may be exposed to this work: We know of
several who provided incorrect advice. Conventional statistical tools designed to deal with
problems related to beer cans coming-off a production line or pesticide treatment on agricultural
plots, may not adequately deal with the special problems of animal population estimation,
such as, unequal sampling probabilities and incomplete detections (Chapter 3).
Many leading quantitative ecologists, mathematicians and statisticians have invested several
decades of first-rate intellectual work to develop the modern approaches to estimation of
animal population parameters we recommend here. Their work bears strongly on what you
do in your tiger surveys. It may not be the wisest use of your time to re-invent these tools.
Your emphasis should be on applying sound techniques to generate good biological data
rather than on inventing new statistical tools. It would be a pity to discard the vast array of
readily available statistical tools and free software in an attempt to do so. (We recommend
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 Monitoring Tiger Populations: Why Use Capture-Recapture Sampling? Monitoring Tiger Populations: Why Use Capture-Recapture Sampling?

that tiger biologists who really believe they have developed a new statistical approach should on camera trap data on tigers, leopards and striped hyenas. Chapters 11 and 12 cover this
submit their innovations to peer review in appropriate journals like Biometrics ). method.

Population estimation models and the software you recommend in this manual appear to be If I want to determine only the relative abundance of tigers from an index, why cannot I use
horribly complicated and difficult to use. simple camera trapping rates (number of individuals caught/100 trap nights) for
comparisons? Why should I still use the capture-recapture sampling framework? Is there a
Not really. Basically, all your field survey work fits in with the following simple canonical dependable relationship between trapping rates and tiger density?
estimator (Equation 3.4) where N is the tiger abundance you are trying to estimate (Chapter 3):
The trapping rate is merely the count statistic C ′ in the above equation divided by the number
C′ of trap-days. If you want to compare trapping rates between two sites as a valid index to the
Nˆ =
pˆ α density that you are trying to estimate, you have to either assume, or make sure, that “p” and
the area sampled are equal between the two areas being compared. Even if you ensure that
where C ′ is the count statistic you obtain in the sampled areas, and pˆ is the estimated detection
the are sampled is equal between two sites, you simply cannot assume that ‘p’s (capture
probability, assumed equal for all samples in this expression (this need not be assumed in
probabilities) are also equal between them. The incorporation of effort (division by number
general; see Thompson 1992). You thus estimate population size N by dividing the count
of trap-days) into the trapping rate index reflects an attempt to remove potential variation in
statistic by both the estimated fraction of the animals on the sampled area that you detected
capture probabilities. However, in addition to effort, these probabilities are influenced by
( pˆ ), and the proportion of the total area from which the count statistic was taken (a ).
trap locations and trap spacing in relation to tiger movement, trail density at the site, and a
host of other unknown and possibly unknowable variables.
If you simply pretend that “p” and “a” are both = 1 in the above equation, they don’t necessarily
become so, and the problem does not go away. On the other hand, you are better off using the
So, it still makes good sense to use a capture-recapture estimation protocol as an integral part
statistical tools that actually deal with the problem of estimating “p” and “a ”, rather than
of your camera trap survey design. If you use capture-recapture sampling, in addition to
pretending that they don’t exist in your world - or assuming without basis that they are both
generating simple trapping rates, you will also get estimates of “p” for both surveys, and you
equal to 1.
can then use trapping rates as a valid index to N if capture probabilities are indeed similar
between sites. Then you will have evidence from your own data to support the claim that ‘p’s
The above sampling-based estimation framework is relevant, if your goal includes any of the
are really equal between the two sites, and your trapping rate index is robust. This material is
four objectives: Estimating the proportion of area occupied (1A), relative abundance (2),
covered in Chapters 7 and 8.
absolute abundance (3), and, survival/recruitment (4). This basic statistical material is covered
in Chapters 3, 4, 7, 9 and 11 of this manual. We note that the above sampling framework has
Let us say you get lucky and catch all the tigers in the area. If you use capture-recapture
even been used for estimating parameters such as species richness, animal movement rates,
sampling, you will have evidence in support of your achievement: The estimated overall
and chances of finding fossils in paleontological surveys, although these aspects are not
capture probability will be close to 1. On the other hand, if you do ad hoc trapping and don’t
covered in this manual.
estimate p at all, you will never have evidence to indicate when you get lucky and catch every
tiger! It would be a real pity if that happened.
Which is the most appropriate sampling-based framework for estimating relative or absolute
abundances of tigers if I decide to use camera traps?
Isn’t capture-recapture sampling more difficult and complicated to implement compared to
the ad hoc schemes I can invent on my own for the camera trap survey?
Capture-recapture sampling is a well-developed method with a sound theoretical basis and
excellent analytical software support. This method allows you to test the fit of probable
The field skills and logistical effort needed for successful camera trapping essentially lie in
alternative estimation models to your own field data (Otis et al. 1978; White et al. 1982;
finding good trap sites and designing a camera placement scheme that ensures that every tiger
Pollock et al. 1990; Burnham and Anderson 1998). We have personally used it successfully
in the sampled area has a chance of getting photographed. Any alternative ad hoc camera-

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 Monitoring Tiger Populations: Why Use Capture-Recapture Sampling?
trapping scheme you invent will also require all the above material resources. There appears
to be no basis for the widely held notion that a capture-recapture scheme is somehow
more complicated and difficult to practice. We argue that the only additional effort
necessary for it is intellectual, involving rigorous survey designs and data analysis. In capture-
recapture surveys you use your time, money, cameras, transport and manpower more
effectively to extract more information from the same field effort. Chapters 11 and 12 cover
this issue.
Should I place the cameras ‘randomly’ within my survey area to sample it properly?
Not using the best trap sites available will lower the capture probabilities (fewer animals
caught and fewer recaptures), thereby weakening your population estimates. Achieving
“randomness” is sometimes said to be the reason for trap placements such as in a square grid
pattern. It should be noted that, as far as possible, sampling should produce similar capture
probabilities for all animals in the sampled area, but the use of a square grid or any other
shape has no particular value in tiger studies. Trap placement that is random with respect to
space will frequently do a poor job of sampling the area over which tigers move.
As explained elsewhere (Karanth and Nichols 1998), ideally there should be no ‘holes’ in the
sampled area within which a tiger with a small home range can have zero probability of
being photographed. Please note that only some tigers will be caught in your sample, but all
tigers present must have some probability >0 of being caught. That is, theoretically, if the
sampling were to be repeated a very large number of times, all animals would be eventually
caught. Chapter 11 deals with this issue.
Placement of traps in areas less likely to be visited by tigers may sometimes be necessary in
order to fill potential holes in a sampled area, but is not otherwise useful for making capture
probabilities equal among individuals or to increase these probabilities. Good trap sites are
difficult to find, particularly in low tiger density areas. Not optimally using best available
trapping sites will decrease data quality. You should optimize trap placement to get more
tiger photos and to increase chances of photographing all the tigers in your sample area.
Is the camera trapping method appropriate for low-density tiger populations?
Like any other method for estimating animal abundance (line transect surveys for example),
the capture-recapture method works relatively better at higher animal densities. Initial
indications from our Indian surveys are that the method yields reasonable results at densities
higher than 2-3-tigers/100 km2 or so. The crucial point is that no other method, including ad
hoc camera trapping, works better at lower densities.
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Monitoring Tiger Populations: Why Use Capture-Recapture Sampling?
Our data suggest that even in low tiger density situations, such as Sundarbans of India (where
we got only six captures, and a density estimate of 0.8-3.6 tigers/100 km2, the capture-
recapture sampling protocol was still amenable to model-based analyses. It was thus a superior
strategy compared to ad hoc trapping. As data and estimates from well-designed camera-
trapping studies become available, it may become possible to “borrow information” from
studies of similar areas in order to get better estimates. There is no reason why (all other
things being equal) a capture-recapture protocol should be more difficult to implement than
ad hoc camera trapping in low-density areas.
On the basis of our studies (Karanth and Nichols 1998, 2000), we do not believe that capture-
recapture sampling requires any greater investment of time, money, or labor than ad hoc
camera trapping. In the worst cases, with poor capture rates or no recaptures, you may still
be able to use the simple removal (Mb) or null models (M0) under a capture-recapture approach.
The Mb model works without recaptures (it is used in analysis of kill-trapping data), although
for tigers Mh is often the preferred model.
Even if all of the closed models fail to work because of sparse and ill-conditioned data, you
still have the option of deriving a naive index based on trapping rates. On the other hand, if
you start with only an ad hoc camera-trapping scheme, you automatically throw away all
chances of performing defensible analyses on the data.
In low-density situations, you may not be able to get reliable estimates of tiger density. At one
such site in India, although tigers were present, we got no captures even after 451 trap nights
of effort. In such cases, you can only hope to get spatial distribution data (Chapters 4, 5 and
6) and should seriously consider whether it is worth investing money and effort into camera
trapping instead of doing simple sign surveys (Chapter 5).
What is the area that I am actually sampling when I do camera trapping?
The sampled area is the area enclosed by all your outermost camera traps, plus a buffer area
around it that includes area within the home ranges of animals sampled by your traps. The
width of the buffer area should be roughly equal to the average half-home range length of
tigers (see Wilson and Anderson 1985; Karanth and Nichols 1998 and Chapter 11 for details).
We are much less comfortable with the idea of computing the sample area as the sum of
arbitrary areas around each single trap point. When each trap site in your scheme is treated as
a distinct ‘sampled area’, it is no longer clear what the sampled population represents, and
indeed you may be sampling several tiger populations at the same time. This issue is covered
in Chapters 11 and 12.
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 Monitoring Tiger Populations: Why Use Capture-Recapture Sampling?
Is there such a thing as camera trapping an area that is too small? How many cameras do
I need to determine tiger density? How large an area do I need to cover? Am I not better off
“over-sampling” rather than “under-sampling” my study area?
Assuming that you want to estimate densities, your goal should be to get as many different
individuals as possible, with the caveat that you want to expose all of the animals in whatever
area you choose, to your sampling effort. By trapping too small an area you may be reducing
the number of individuals that are potentially exposed to your camera traps. By increasing
trapping effort over a smaller area, you increase chances of recaptures of the same individuals
leading to better estimates of capture probabilities. If you have a limited number of cameras,
as you expand the area to catch more individuals, the recapture rates will drop. Therefore,
there is a trade-off between increasing the sample size (the number of individual tigers caught)
on the one hand, and, ensuring no “holes” in the study area and increasing capture probabilities
of fewer individuals on the other.
The reliability of your density estimate depends both on the number of tigers captured and the
estimated capture probabilities. There is no “safe” rule of thumb, such as “over-sampling”
being better than “under-sampling”, etc., for getting your trap spacing right. It really depends
on the logistical constraints you face (time available, number of cameras, ability to cover the
area) and your best guess of what a breeding female’s range size may be in the area.
You should place your camera traps so that a resident (breeding) tigress is exposed to at least
2-3 cameras given the expected home range size of such females in the area. Since the smallest
recorded ranges in the high productivity habitats in south Asia are around 10-15 km2 or so
(where one can safely use a 2-3 km-apart trap spacing), in areas with lower density (larger
female ranges) much greater trap spacing can be used “safely”. If we try to be too “safe” by
putting cameras very close to each other, we reduce the area sampled and the total number of
different individuals potentially exposed to our traps. This issue is covered in Chapters 11
and 12.
Which capture-recapture models are most appropriate for estimating tiger abundance?
An unjustified assumption, often implicitly made by some tiger biologists, is that all individual
tigers in the sampled area have the same probability of being photo-captured. This is
very unlikely. Placing your traps in some sort of a grid pattern, to obtain what is believed
to be “a random sample” (see White et al. 1982; Thompson et al. 1998 for details) does
not make the problem of unequal catchability go away. As noted above, a trap placement
that is random with respect to space frequently does a poor job of sampling the locations
and movements of tigers within that space. On the other hand, using model Mh or another
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Monitoring Tiger Populations: Why Use Capture-Recapture Sampling?
model incorporating heterogeneity explicitly deals with this problem (Otis et al. 1978).
Capture probabilities vary due to factors such as trap location in relation to home range and
movements of each individual and its social position (residents, transients etc.). The
probabilities may (although less likely) vary due to trap response, time of trapping, etc. Your
essential challenge is to estimate these variable capture probabilities using a model that
approximates the process most likely to have generated the capture histories you got. Computer
programs like CAPTURE and MARK provide you with model selection tools to help you
select a reasonable approximating model. The paucity of capture-recapture data in tiger
studies may lead to selection of simple models (e.g., M0), and these models may provide
reasonable estimates of densities. We have tended to favor use of Mh because of its robustness
when assumptions are not met, but this simply reflects personal judgment and should not be
viewed as any sort of rule.
Why should I use two cameras per trapping point as opposed to only one? The latter is a
cheaper option and allows me to cover a larger area and get more captures.
You can use a single camera only if you can have the camera placed above the tiger’s path
photographing the stripe patterns on the tiger’s back. Of course, you will lose information on
the age-sex class of the tiger when you do so. Also, the pictures are not very pretty to show
around! If you use flanks or frontal shots of tigers for identifications, you need two cameras.
Making unjustified assumptions based on single flank pictures (this tiger was photographed
here so must be different from that tiger photographed there, etc.) is not good science and
may lead to incorrect inferences. Tigers move long distances, and such assumptions will
leave you open to criticism. While it may be possible to develop statistical models that can
include the additional uncertainty of using single flank photos, such models are not available
at present. When they do become available, resulting estimates will have larger variances
reflecting the uncertainty associated with individual identification. Other than telling stories
about your tigers, there is no way of getting around taking pictures of both flanks.
How long should I camera trap to get reliable results? Weeks, months, years?
This is a compromise between the area you want to sample, the number of traps you have and
how intensively you want to camera trap. But you cannot extend the sampling duration
indefinitely: A critical assumption of closed models is that tigers are not moving in or out of
the sampled population and that no recruitment and deaths occur during your sampling. The
shorter the sampling duration, the better it is for meeting the closure assumption. However,
tigers being fairly long-lived animals, we can perhaps survey for a few weeks and still assume
population closure.
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 Monitoring Tiger Populations: Why Use Capture-Recapture Sampling?
If the closure assumption is violated, then models for open populations can still be used with
your data. Such models for open populations are implemented in programs such as POPAN,
JOLLY, JOLLYAGE, and these programs can be used for estimating population size. These
open models are not as versatile as the closed models implemented in programs CAPTURE
and MARK for estimating population size, in the sense that they cannot be readily adapted to
deal with some sources of variation in capture probability.
Where do I use open models? And how?
Open models are primarily useful for estimating survival and recruitment from capture-
recapture data obtained by doing annual surveys at the same site, year after year. In such
long-term studies of tigers, you can use the closed models for estimating abundance each
year, and open models for estimating survival rates between years. Recruitment is then estimated
using a combination of models for open and closed populations. The two types of models can
be combined in what is called the Robust Design (Pollock et al. 1990). Under the robust
design, estimation can either be done in separate steps using first the closed and then open
models (e.g., Pollock et al. 1990), or a joint model can be developed to deal with all estimation
at one time (Kendall et al. 1995, 1997). Program MARK (White and Burnham 1999) can be
used with either approach.
Can I use individual identifications of tigers based on shapes of their track prints in capture-
recapture surveys?
The statistical track shape discrimination methods have so far worked only on small captive
populations (N<12-17) in tests under controlled conditions (Riordan 1998). However,
unambiguous identifications cannot be derived easily in the field. Variations in substrate
conditions, slope of terrain and speed of the animal, and similarities between tiger tracks, all
combine to create ‘different animals’ for the track discrimination software. The software is
then ‘fooled’ into giving you ‘tiger identifications’ that are in fact merely the number of
‘different looking’ track shapes that the software’s algorithms recognize. Further, these
identifications are probabilistic (this track could be of tiger X with a probability of 0.6). Such
probabilistic identifications cannot be used in currently available capture-recapture models.
If I camera trap tigers, should I also do radio telemetry? Can I use camera trap captures for
home range estimation instead of radio telemetry?
Radio telemetry gives movement data that can be incorporated into better estimation of the
sampled area (Chapters 11 and 12). Accounts of innovative use of telemetry data in combination
with capture recapture methods have been published recently (Powell et al. 2000). See if you
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Monitoring Tiger Populations: Why Use Capture-Recapture Sampling?
can use telemetry data to better estimate movement and range size. But telemetry is expensive
and tigers are very difficult to catch and radio-track!
In practice, camera trap photos do not yield enough data points for reliable home range
analysis. Moreover, unlike telemetry locations, which are a representative sample of all possible
locations that a tiger visits in its home range, camera trap captures represent only some fixed
points within the animal’s home range. These points are not a representative sample of the
animal’s home range use. They can only give a rough idea of home range size and movement
distances.
I don’t seem to be getting tiger cubs in my camera traps. Some tigers seem to be avoiding
my traps. What can I do about these problems?
Yes, cubs less than a year old do seem to have much lower capture probabilities, and it may
take too much effort to estimate their numbers from capture-recapture surveys. In a sense,
they are like a different species with inherently much lower capture probabilities. One can
guess at the numbers of cubs as a proportion of the total population based on the number of
breeding females in the area, or from demographic models of tiger populations (Karanth and
Stith 1999).
If deliberate trap avoidance occurs on a significant scale in a given data set, program
CAPTURE will detect that, and model selection results will indicate use of models that
incorporate effects of trap response (Mb or Mbh models), as these are more appropriate for the
analysis of such data. More details are provided in Otis et al. (1978) and White et al. (1982)
and Chapter 11.
Can camera trap photos of prey species give me information on their numbers?
Unless the individuals of a prey species are uniquely marked (as in chital deer), we cannot
derive capture-recapture estimates of prey abundance. Even if some prey species are uniquely
marked, the trap spacing appropriate for sampling prey and tiger populations will be different.
The best you can hope with prey species is a trapping rate index of density (number of
sambar photos/100 trap nights). If a camera trap happens to be set in a particular sambar’s
favored saltlick, the same animal will be caught several times. In such a case, the trapping
rate index may not have much of a relationship to true sambar density. However, such trapping
rate indices of prey abundance are often ‘by-products’ of surveys for tigers. If at all possible,
try to get independent prey density estimates from line transect surveys or at least dung
counts (Chapters 7, 8, 9 and 10).
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 Monitoring Tiger Populations: Why Use Capture-Recapture Sampling?
I have two choices: I can try to estimate tiger densities at a few sites using the rigorous
methods you suggest or I can go out and generate ‘quick and dirty’ estimates at many more
places in the same region? Given the scarcity of data on wild tigers, which approach is
more useful for conservation?
Our data (Karanth and Nichols 1998, 2000) suggest that tiger densities can vary between
0-20 animals/100 km2 over the vast range of the species. At any given site that you choose,
the possible value for tiger density will fall within an even narrower range. Therefore, if we
get reliable estimates from some representative sites across the geographic range of tigers,
we can make informed guesses about many other areas based on our knowledge of tiger
biology and conservation issues. Therefore, we can accomplish many conservation goals
based on such reliable estimates. So, right now the critical conservation need is to get reliable,
scientifically defensible estimates of tiger densities from at least a few sites. It is far more
useful to have a “few” reliable tiger density estimates, rather than possess “many” unreliable
“density estimates of some kind”. In fact, using such ‘quick and dirty’ tiger abundance estimates
derived from many sites, wildlife managers are already making serious conservation errors
that the endangered big cat can ill-afford (Karanth et al. in prep.).
REFERENCES
Buckland, S.T., Anderson, D.R., Burnham, K.P. and Laake, J.L. 1993. Distance sampling:
Estimating abundance of biological populations. Chapman and Hall, London, UK.
Burnham, K.P., and Anderson, D.R. 1998. Model selection and inference: A practical
information theoretic approach. Springer-Verlag, New York, NY, USA.
Johnson, D.H. 1999. The insignificance of statistical significance testing. Journal of Wildlife
Management 63:763-772.
Karanth, K.U. and Nichols, J.D. 1998. Estimating tiger densities in India from camera trap
data using photographic captures and recaptures. Ecology 79:2852-2862.
Karanth, K.U. and Nichols, J.D. 2000. Ecological status and conservation of tigers in India.
Final Technical Report to the US Fish and Wildlife Service (Division of International
Conservation), Washington, DC, and Wildlife Conservation Society, New York. Centre
for Wildlife Studies, Bangalore, India.
Karanth, K.U. and Stith, M.B. 1999. Prey depletion as a critical determinant of tiger population
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Monitoring Tiger Populations: Why Use Capture-Recapture Sampling?
viability. Pages 100-113 in Riding the tiger: Tiger conservation in human-dominated
landscapes (Eds: J. Seidensticker, S. Christie and P. Jackson). Cambridge University
Press, Cambridge, UK.
Karanth, K.U., Nichols, J.D., Seidensticker, J., Dinerstein E., Smith, J.L.D., McDougal C.,
Johnsingh, A.J.T., Chundawat, R.S. and Thapar, V. Science deficiency in conservation
practice: The monitoring of tiger populations in India. (In Prep.).
Kendall, W.L., Pollock, K.H. and Brownie, C. 1995. A likelihood-based approach to capture-
recapture estimation of demographic parameters under the robust design. Biometrics
51:293-308.
Kendall, W.L., Nichols, J.D. and Hines, J.E. 1997. Estimating temporary emigration using
capture-recapture data with Pollock’s robust design. Ecology 78:563-578.
Lancia, R.A., Nichols, J.D. and Pollock, K.N. 1994. Estimation of number of animals in
wildlife populations. Pages 215-253 in Research and management techniques for
wildlife and habitats. (Ed: T. Bookhout). The Wildlife Society, Bethesda, MD, USA.
Nichols, J.D. 1992. Capture-Recapture models: Using marked animals to study population
dynamics. Bioscience 42:94-102.
Otis, D.L., Burnham, K.P., White, G.C. and Anderson, D.R. 1978. Statistical inference from
capture data on closed animal populations. Wildlife Monographs 62:1-135.
Pollock, K.H., Nichols, J.D., Brownie, C. and Hines, J.E. 1990. Statistical inference for
capture-recapture experiments. Wildlife Monographs 107:1-97.
Powell, L.A., Conroy, M.J., Hines, J.E., Nichols, J.D. and Krementz, D.G. 2000. Simultaneous
use of mark-recapture and radio telemetry to estimate survival, movement, and capture
rates. Journal of Wildlife Management 64:302-313.
Riordan, P. 1998. Unsupervised recognition of individual tigers and snow leopards from their
footprints. Animal Conservation 1:253-262.
Thompson, S.K. 1992. Sampling. Wiley, New York, NY, USA.
Thompson, W.L., White, G.C. and Gowan, C. 1998. Monitoring vertebrate populations.
Academic Press, New York, NY, USA.
165
 Monitoring Tiger Populations: Why Use Capture-Recapture Sampling?

White, G.C. and Burnham, K.P. 1999. Program MARK: Survival rate estimation from both APPENDIX 1.1
live and dead encounters. Bird Study 46:120-139.
DRAWINGS OF THE TIGER AND OTHER SYMPATRIC LARGE CARNIVORES AND THEIR
White, G.C., Anderson, D.R., Burnham, K.P. and Otis, D.L. 1982. Capture-recapture and TRACKS
removal methods for sampling closed populations. Los Alamos National Laboratory
Publication LA-8787-NERP. Los Alamos, NM, USA. Front Foot

Williams, B.K., Nichols, J.D. and Conroy, M.J. 2002. Analysis and management of animal
populations. Academic Press, San Diego, CA, USA.

Wilson, K.R., and Anderson, D.R. 1985. Evaluation of two density estimators of small mammal
population size. Journal of Mammalogy 66:13-21.

Yoccoz, N.G., Nichols, J.D. and Boulinier, T. 2001. Monitoring of biological diversity in Tiger
space and time. Trends in Ecology and Evolution 16:446-453.

Front Foot

Asiatic Lion

Front Foot

Leopard

Front Foot

Snow Leopard

166 Drawings not to Scale

167
 Appendix 1.1 Appendix 1.1

Front Foot Hind Foot Front Foot

Hyena

Asiatic Black Bear

Front Foot Hind Foot

Front Foot

Wolf

Sloth Bear

Front Foot Hind Foot

Front Foot

Malayan Sun Bear Dhole

168 Drawings not to Scale Drawings not to Scale

169
 Appendix 1.2

APPENDIX 1.2

DRAWINGS OF PRINCIPAL PREY SPECIES OF THE TIGER AND THEIR TRACKS

Front Foot Front Foot

Elephant Calf
Gaur

Hind Foot

Front Foot

Rhino Calf

Banteng
Front Foot

Asiatic Water Buffalo

170 Drawings not to Scale Drawings not to Scale

171
 Appendix 1.2 Appendix 1.2

Front Foot Hind Foot

Front Foot

Malayan Tapir

Barasingha
Front Foot

Front Foot

Nilgai

Eld’s Deer

Front Foot

Sambar

172 Drawings not to Scale Drawings not to Scale

173
 Appendix 1.2 Appendix 1.2

Front Foot

Front Foot

Hog Deer

Chital
Front Foot

Front Foot

Muntjac

Hard Soil
Wild Pig

Soft Soil

174 Drawings not to Scale Drawings not to Scale

175
 Appendix 1.3

APPENDIX 1.3 observation. If you are unable to figure out the position of the limb as described above, record
it as NA. Examples: TT-LH, TT-NA
SPECIMEN DATA FORM FOR FIELD SURVEYS OF SPATIAL DISTRIBUTION OF TIGERS
Specimen Reference: If the survey team collected a specimen (plaster cast, scat) they found
AND PREY
and recorded it in the previous column, give it a unique specimen reference number. Link the
Form No: Reference Map Number: Date of Survey: Leader: specimen number to the sample data form number and the date of survey. For example: The
first tiger pugmark plaster cast specimen collected during sample survey number 22 in
Number of Personnel: Start Time: Stop Time: Bandipur on 6-6-2001 will be recorded as TC-1, the third tiger scat found on the same survey
Survey Effort (Hours): as SC-3, and so on in this column. Make sure that you attach the specimen reference number
on a label using indelible ink to all specimens. For example: TC-1/Bandipur-22/6-6-2001.
Name/Identity of the Area Surveyed:
Approximate Latitude and Longitude of the Surveyed Area: Locality: Ideally this location should be obtained in latitude and longitude with a GPS unit.
However, if GPS units are not available, describe in detail the locality where the sighting of
Approximate Distances covered by the Survey Team during the day (Km):
animal or animal sign occurred to the nearest 100 or 1000 meters accuracy.
Walking: Vehicle: Boat: Elephant Back: Other Transport: Total:
Map ref: Map number and the ID number of the location related to this observation marked
Detailed Description of the Route and Names of Localities Visited by the Survey team: on the map: Example: MR 13/Bandipur22/Bandipur Map-6.

In the following data table, if information was not recorded write NR; if not observed accurately
or not available write NA; if not applicable write NP; and if no specimen was collected write Data Table
NC.
Sl. Specimen
S. No.: This is the serial number of that particular observation\data point Time Species Nos. Specimen Locality Map Ref.
No. Reference
Time: Time of making the observation: e.g., 5.30 PM or 1730 hours

Species: Animal species seen or signs detected; e.g., Sambar, Chital etc. 1

Nos.: How many animals were seen? If sightings or tracks of tigers accompanied by cubs
are detected, indicate this. For example, if tracks of a tigress with 2 cubs were seen, record
this as: 1F+2C; If a male and female tiger were seen record as: 1F+1M and so on. If you were
2
not able to record age-sex etc., record only the numbers e.g.: 1. If you could only count
approximately (as is often the case with prey species), record as approximate numbers: 1-5;
30-40 or > 50 etc., and so on. If you were unable to count the animals at all, record the
number as NA.
3
Specimen: Record what type of specimen was observed or collected (track plaster cast = TC;
track tracing = TT, scat = SC, prey dung = PD, kill or part of kill = KL). If you see a set of
tracks and follow them over long distances, record the detection only once. Do not repeatedly
record the same set of tiger tracks. If imprints of all four paws are collected, identify the 4
specimens as: Right Front (RF), Left Front (LF), Right Hind (RH) and Left Hind (LH). In
this case there will be four specimens and four specimen reference numbers for a particular

176 177
 Appendix 1.4

APPENDIX 1.4 6. Occurrence of tiger poaching in the surveyed area in the past one year:
Organized tiger poaching or tiger trade (Yes / No / No information)
SPECIMEN DATA FORM FOR QUESTIONNAIRE SURVEYS OF INFORMANTS Incidental tiger poaching (Yes / No / No information)

Form No: 7. Occurrence of hunting or poaching of prey species:

Date of filling the form: Heavy Moderate Occasional No Information

The area to which survey data in the form pertains:

8. Details of any man-killing or man-eating by tigers in the surveyed area durin past one
Reference map linked to the data form: year:

Name and address of the person filling the form:

9. Non Timber Forest Produce collection impact: Severe Moderate No No information
Informant’s name and contact address:
Basis for the data provided: Field Visits/Secondary Informants/Both 10. Wood\Biomass extraction (legal): Severe Moderate No No information

If secondary data (reports, hearsay etc.) are used, details: 11. Wood\Biomass extraction (illegal): Severe Moderate No No information

1. Assessment of tiger population trend during last five years within the survey area: 13. Livestock grazing impact: Severe Moderate No No information
Stable Declining Increasing No information
14. Forest fire impact: Severe Moderate No No information
2. Evidence for tiger presence during past one year (‘how do you know tigers are present
now?’): 15. Impact of developmental projects either planned or underway:
Tigers seen Tiger signs seen Secondary Source No evidence Highways: Yes No No information
If the informant can give details, mark the locations where tigers are present on the reference Local roads: Yes No No information
map and record the map reference:
Mines: Yes No No information
3. Evidence for Tigers reproducing (‘how do you know there are cubs being born?’) Dams: Yes No No information
Cubs Seen Tracks seen Multiple Kills Secondary Source No evidence Canal: Yes No No information
If the informant can give details, mark the locations where tigers are reproducing on the Industrial plants: Yes No No information
reference map and record the map reference: Agricultural conversion: Yes No No information

4. Assessment of prey base: Name and rank prey species in order of importance (including Tourism: Yes No No information
livestock). Others (please specify): Yes No No information
1. 2. 3. 4.
16. Any other relevant anecdotal information:
5. Presence of other predator species, within tiger habitat in the survey area
Leopard Snow Leopard Dhole Hyena Wolf

178 179
 APPENDIX 2.1 APPENDIX 2.2

SPECIMEN DATA FORM FOR ENCOUNTER RATE SURVEYS OF TIGER SCATS SPECIMEN DATA FORM FOR RECORDING DUNG COUNTS ON PLOTS

Field Site: Page No.:

Biologist: Field Site:
Biologist: Start Location:
Date: Weather: Page No.: Date: End Location:
Start Time: End Time: Sample Walk No.:
Plot Plot No. of Pellet Piles in each Decay Stage Habitat
Start Location: End Location: Species
No. Location A B C D Type
Predator
Evidence Scat
Sl. Specimen Time of (Tiger,
Location (Size/Scrape/ Diameter Prey ID
No. No. Collection Leopard, Total
Tracks) (in mm.)
Unknown)

Total

Total

Total

Total

Total

Total
DECAY STAGES:
A: Fresh and moist with odor. B: Completely dry with inner contents also completely dry. C: Decomposition
has begun; a few pellets have started disintegrating. D: 50% or more of the pile is decomposed, but still
recognizable as a unique pile of a given species. E: Totally decomposed and unidentifiable.
HABITAT TYPE:
BSC: Bare soil. OPN: Short grass clearing with open canopy.
CCP: Moderate to dense undergrowth with closed canopy.

180 181
 APPENDIX 3.0 APPENDIX 4.0

SPECIMEN DATA FORM FOR LINE TRANSECT SURVEYS SPECIMEN DATA FORM FOR CAMERA TRAP SURVEY OF TIGERS

Biologist: Transect Line No.: Field Site: Field Site: Data Form No:
Date: Weather: Page No.: Biologist: Sample No.:
Start Time: End Time: Sample No.: Exposure details Events
Unit
Start Location: End Location: Date Time Location Cam Roll Remarks
No. ST CL EXPS ST CL
No. No.
Sighting Compass Bearings Perpendicular
Nos. Habitat/ Setup
Time Species Distance Animal Line Angle Distance
s Location
r θ1 θ2 q x
Check

Setup

Check

Setup

Check

Setup

Check

Setup

Check

Unit No. = Camera Trap Equipment ID number Events

Cam No. = Camera ID number ST = Starting event number
Roll No. = Film roll ID number CL = Closing event number
ST = Starting frame number of the Film roll
CL = Closing frame number of the Film roll
EXPS = Number of exposures during the trap period

182 183
 Appendix 5.0

APPENDIX 5.0 JAVAJI METAL SECURE CAMERA TRAP SHELL

DESIGN OF THE JAVAJI METAL SECURE CAMERA TRAP SHELL

A number of commercial manufacturers are now making camera-traps. Many biologists

and wildlife managers using such expensive camera traps in field surveys face major problems
of theft, tampering by humans and animal damage. The Javaji Metal Secure Camera
Trap Shell is a protective device, which is designed to reduce the risk of theft, tampering and
animal damage to the camera traps. The illustrations on the next page depict the Javaji Metal
Secure Shell designed for use with Trailmaster, which is one of the leading brands of
camera-traps in the market. With modifications, it can be used to protect other makes of
camera-traps also.

The shell has been designed and developed on a non-commercial basis by engineer-
conservationist Javaji Amarnath, Bangalore, India. It was developed under the guidance of
K.U. Karanth. The fabrication and field testing of the shell was done in collaboration with
N. Samba Kumar, G.R. Sanath Kumar and Jeevan Rao. The final technical drawings
Front View Rear View
were prepared by Niren Jain.

We intend to share this design freely with other biologists. However, we provide no guarantees
to any user about its efficacy or effectiveness. Any use of this design or unit will be at the
risk of the user, and its developers are not responsible for any losses or damage resulting
from its use. Additional information on the fabrication and use of this shell can be obtained Transparent Camera Box
Camera Box
Antiglare Shield
from the WCS India Program (Email: <wcsind@bgl.vsnl.net.in>) in the form of Adobe
Acrobat Reader pdf file.
Washer
Sleeve
Pivot Axle

Camera Box Housing

Pivot Axle

Camera box pivot axle assembly

184 185
 Appendix 5.0

APPENDIX 6.0
Locking
Link WEB SITES FOR FREE ANALYTICAL SOFTWARE AND ADDRESSES OF SOME CAMERA
TRAP EQUIPMENT SUPPLIERS
Chain
Locking
Housing
Pin Web Sites for Free Software
Chain
Locking The program CAPTURE is downloadable from the web at the sites of the USGS Patuxent
Plate
Wildlife Research Center, Laurel, Maryland. This site is maintained by Jim Hines.
Lock Bolt http://www.mbr-pwrc.usgs.gov/software.html
Lock Nut
Allen Screw of
The software CAPTURE, MARK, JOLLY, JOLLYAGE, etc. and the out of print Wildlife
Chain Chain Locking Assembly Housing Collar Monograph of Otis et al. (1978: Chapter 11) referred to above, are available from the Web
Site of Gary White at Colorado State University, Fort Collins, Colorado. This site is maintained
by Gary White who also maintains the MARK list server discussion group at the Colorado
Extension site that deals with capture-recapture issues.
Coloumn
http://www.cnr.colostate.edu/~gwhite/mark/mark.htm

The programs DISTANCE 3.5 and DISTANCE 4.0 BETA 3 and the out of print book
Buckland et al (1993: Chapter 9) are downloadable from the website of Centre for Research
Locking Plate Holding Tool
into Ecological and Environmental Modelling (CREEM). This site is maintained by Len
Thomas at the Research Unit for Wildlife Population Assessment, University of St. Andrews,
Pedestal
Scotland, UK. http://www.ruwpa.st-and.ac.uk/distance/

Camera Trap Equipment Suppliers

D-Shackle Buckshot 35 mm Game Scouting Camera
http://www.buckshot35.com/buckshot35a.htm
D-Shackle Pin

Cross Bar CamTrakker – The big Buck Surveillance system

Ground Anchor
http://www.camtrakker.com/

Gametronix GameScout
Ground Anchor
http://www.gametronix.com/home.html
Exploded view
Game-Vu – Digital trail camera system
http://www.gamevu.com/

Locking the chain to the ground anchor Highlander Photoscout

http://www.highlandersports.com/

186 187
 Appendix 6.0

Moultrie Got-cha APPENDIX 7.0

http://www.moultriefeeders.com/
GLOSSARY OF TECHNICAL TERMS
Phantom Hunter
http://www.phantomhunter.com/ Adapted from Thompson, W.L., White, G.C. and Gowan, C. 1998. Monitoring vertebrate
populations. Academic Press, New York, NY, USA.
TrailMAC
http://www.trailsenseengineering.com/
Abundance: Total number of individuals or items of interest in some defined area and time
Trailmaster – Trail monitoring systems period; also known as absolute abundance.
http://www.trailmaster.com/
Bias: A persistent statistical error associated with parameter estimates whose source is not
Trail Timer – Game Monitors random chance. More precisely, bias is the difference between the expected value of a parameter
http://www.trailtimer.com/ estimate and the true value of the parameter. For example, a negatively biased estimator
produces estimates that, on average, are smaller than the true quantity being estimated.
Vigil -Trail Infrared Monitor
http://www.roc-import.com/gb/monitor/vigil_gb.html Census: A complete count of individuals, objects, or items within a specified area and time
period. A census generally refers to a complete count of all elements within a sampled
WoodsWatcher population and/or target population; however this term also may be applied at the level of the
http://www.woodswatcher.com/ sampling unit to represent a complete count of elements within a sampling unit, such as a
“plot census”. “Census” is frequently misused as a synonym for “survey”. True censuses are
extremely rare in work with wildlife populations.
For a review and comparison of the performances and specifications of these camera trapping
units, visit this website: http://www.jesseshuntingpage.com/cams.html Closed population: A fixed group of individuals within a defined area and time period, i.e.,
there are no births, deaths, immigration and emigration on the area for the period of
interest.

Coefficient of variation (CV): Ratio of a standard error of a parameter estimate to the

parameter estimate. The coefficient of variation is used in computing sample sizes and as a
measure of relative precision when comparing degree of variation among different estimates
or sets of data.

Confidence interval (CI): An interval around a parameter estimate that provides a measure
of confidence regarding how close a sampled-based estimate is to the true parameter. The
usual two-sided symmetrical confidence interval around the parameter estimate is generated
by adding and subtracting the quantity computed from the product of the standard error and
the t value or z value corresponding to the pre-specified (1-K) % confidence level (K is
frequently set at 0.05). For example, a 95% confidence interval will contain on average, the
true parameter of interest 95 of 100 times if 100 such intervals were calculated in a like
manner. That is, confidence refers to the procedure of obtaining an interval rather than the

188 189
 Appendix 7.0
interval itself. There is not a 95% probability that the true parameter occurs in the interval;
either a parameter is in the interval or it is not.
Covariate: A variable that may be related to a parameter of interest. Sometimes this relationship
is of direct interest, as when tiger abundance is modeled as a function of prey density, where
prey density is viewed as the covariate. In other cases, the covariate relationship is not of
intrinsic interest itself. In an analysis of covariance, the relationship between the dependent
variable and the covariate is first adjusted for before the effects of the other factors are
examined.
Density: Total number of individuals or objects of interest per unit area (also known as
absolute density). Sometimes, the concept is broadened to mean number of animals per unit
resource, where resource could be suitable habitat, food abundance, etc.
Detectability: Probability of correctly noting the presence of an element within some specified
area and time period. Detectability can also be viewed as the expected proportion of elements
that is detected.
Estimate: A numerical value calculated from sample data collected from a sampled population
and used to represent the parameter of interest.
Estimator: A mathematical formula used to calculate an estimate. An unbiased estimator
will produce unbiased estimates when appropriate assumptions are satisfied.
Index: A relative measure used as an indicator of the true state of nature.
Index of relative abundance: Count statistic believed to provide a proportional measure of
the number of individuals within an area.
Index of relative density: Count statistic believed to provide a proportional measure of the
number of individuals per unit area.
Georeference: Georeferencing establishes the relationship between an image (row, column)
coordinate system and a map (X,Y) coordinate system. X, Y map coordinates are translated
into latitude/longitude real world coordinates which point to geographic features, such as
street names and addresses. Also known as geocoding.
Monotonic relationship: A relationship that is continually increasing (or decreasing).
190
Appendix 7.0
Open population: A group of individuals whose number and composition are not fixed within
a defined area over a period of interest, i.e., there could be births, immigration, deaths, and/
or emigration on the area over the period of interest.
Parameter: An unknown numerical quantity or constant associated with some measure of a
target population.
Plot: A sampling unit of some defined area or volume.
Population trend: An average change over time in magnitude and direction of some population
parameter within a specified area.
Precision: The degree of spread in estimates generated from repeated samples. Variance,
standard deviation and standard error are all measures of precision.
Probability distribution: The probability structure generated from all possible values of
some random variable.
Random sample: A collection of sampling units chosen based on known chance of selection.
Random selection allows some probability or degree of certainty to be attached to resulting
sample estimates in order to assess their usefulness.
Robustness: The ability of an estimator to produce estimates with relatively small bias even
if the underlying assumptions are not met.
Sample: A group of sampling units selected during a survey.
Sampled population: All elements associated with sampling units listed or mapped within
the sampling frame.
Sampling design: Protocol for obtaining parameter estimates for a sampled population. The
purpose of a sampling design is to make inferences to the sampled population, usually in
conjunction with an observational study. In monitoring, a spatial sampling design specifies a
means of selecting spatial sampling units that permits inference about the sampled population.
A sampling design may also be used in an experimental approach for obtaining estimates of
differences in parameters associated with treatment groups.
Sampling distribution: The probability distribution of a sample estimate based on probability
of occurrence of estimates generated by all possible samples of a given size.
191
 Appendix 7.0

Sampling unit: A unique set usually of one or more elements. In spatial or area sampling a APPENDIX 8.0
sampling unit (e.g., plot of ground) may not contain any elements.
COMMON AND SCIENTIFIC NAMES OF ANIMALS USED IN THE TEXT

Sampling variation: A measure of the degree of spread whose source is solely from random
chance associated with the selection procedure (i.e., among-unit variation) and/or counting Hanuman Langur - Semnopithecus entellus
protocol (i.e., enumeration variation). Wolf – Canis lupus
Dhole – Cuon alpinus
Spatial distribution: A geographical range of locations or areas occupied by a species. Malayan Sun Bear – Ursus malayanus
Sloth Bear – Ursus ursinus
Standard deviation: Square root of variance of individual items in a probability distribution.
Asiatic Black Bear – Ursus thibetanus
In this case, “distribution” refers to either the true or population distribution, such as the
distribution of all plot abundances, Ni (called the population standard deviation), or the Hyena – Hyaena hyaena
distribution within a single sample, such as the distribution of items within a single plot Asiatic Lion – Panthera leo
sample (called the sample standard deviation or just “standard deviation”). Leopard – Panthera pardus
Tiger – Panthera tigris
Standard error: Square root of variance; the standard deviation of a sampling distribution
Snow Leopard – Uncia uncia
of sample estimates. The “population standard error” describes this measure for a sampling
distribution of all possible sample estimates. An estimator of this quantity, called the sample Asian Elephant – Elephas maximus
standard error (or just “standard error”), may be obtained from a single sample and, for Malayan Tapir – Tapirus indicus
infinite populations, is equal to the sample standard deviation divided by the square root of Greater One-horned Rhinoceros – Rhinoceros unicornis
sample size. The standard error is especially useful for computing a confidence interval for a Wild Pig – Sus scrofa
parameter estimate.
Chital – Axis axis
Hog Deer – Axis porcinus
Survey: A count (usually incomplete) of individuals, objects or items within a specific area
and time period. Eld’s Deer – Cervus eldii
Sambar – Cervus unicolor
Trend: A change in average status of some quantity or attribute over a defined time period. Barasingha – Cervus duvaucelii
Muntjac – Muntiacus muntjak
Variance: A measure of precision; average of squared differences between a set of values and
Banteng – Bos javanicus
the mean of the distribution of those values.
Gaur – Bos gaurus
Nilgai – Boselaphus tragocamelus
Asiatic Water Buffalo – Bubalus bubalis

Source:

Corbet, G.B. and Hill, J.E. 1992. The mammals of the Indo-Malayan region: A systematic
review. Oxford University Press, London, UK.

192 193