Aspects of Cheetah (Acinonyx jubatus) Biology, Ecology and

516 Pages·2003·4.8 MB·English

Aspects of Cheetah (Acinonyx jubatus)
Biology, Ecology and Conservation Strategies on
Namibian Farmlands

by
Laurie L. Marker

Thesis submitted for the degree of
Doctor Of Philosophy

Lady Margaret Hall, University of Oxford
Trinity Term 2002
ASPECTS OF CHEETAH (ACINONYX JUBATUS) BIOLOGY, ECOLOGY AND
CONSERVATION STRATEGIES ON NAMIBIAN FARMLANDS

Summary
In an increasingly human-dominated environment, the task of successfully conserving
large carnivores, such as cheetahs, is difficult due to real or perceived threats resulting in conflict
and often their local extirpation. This research describes the causes and potential solutions to this
conflict in Namibia. Cheetah biology and ecology were studied through physical examination,
laboratory analysis, radio-tracking and human perceptions using survey techniques.
Between 1991 and 2000 data collected on over 400 live-captured and dead cheetahs
showed that a perceived threat to livestock or game was the reason for 91.2% (n = 343) of
cheetahs captured and 47.6% (n = 30) of wild cheetah deaths. Both were biased towards males,
with 2.9 males being captured for every female, despite an apparent equality of sex ratio.
Human-mediated mortality accounted for 79.4% (n = 50) of wild deaths reported, of which the
majority involved prime adult animals, with a peak at around 5-6 years of age.
Polymorphic microsatellite loci were used to assess 313 Namibian cheetahs’ variation,
gene flow, paternity and behavioural ecology. Genetic analysis showed limited regional
differentiation supporting a panmictic population and that persistence in Namibia depends on
dispersal from regions throughout the country; therefore efforts of connectivity throughout the
country should continue. Relatedness values confirmed family groups, and 45 new potential
sire/dam offspring and 7 sibling groups were identified, providing information on dispersal and
the success of translocation. Sera from wild cheetah were assessed for exposure to feline and
canine virus antibodies to CDV, FCoV/FIP, FHV1, FPV, and FCV; antibodies were detected in
24%, 29%, 12%, 48%, and 65%, respectively, showing infection occurs in wild cheetahs;
although there was no evidence of disease at time of capture, these diseases are known to cause
serious clinical disease in captive cheetahs. Neither FIV antibodies nor FeLV antigens were
present in any wild cheetahs tested, however, the first case of FeLV in a non-domestic felid is
described in a captive Namibian cheetah. Concern for contact with domestic animals is
discussed. Focal Palatine Erosion (FPE), a dental abnormality found in captive cheetahs, was
discovered in over 70% of the wild cheetahs and was correlated with dental malocclusions, and is
of concern to the long-term health of wild cheetahs.
Namibian cheetahs have a mean 95% kernel home range of 1642.3 km2 (+ 1565.1 km2),
the largest home ranges yet defined. Habitat type significantly affected the cheetah’s spatial
distribution and prey density. Radio-collared female cheetahs were more closely related to other
cheetahs in the study area than males, indicating male dispersal. Continual cheetah perturbation
may partially explain the unusually low density of cheetahs in this area (estimated at only 2.5
cheetahs per 1000km2) despite the apparent abundance of prey.
Namibian farmers originally surveyed revealed a mean removal of 19 cheetahs per
year/farm, even when not considered a problem, and higher removals occurred on game farms.
Evidence for actual livestock depredation was negligible, only 3% of reported captures. Scat
analysis revealed cheetahs’ selection for indigenous game, however 5% of scats contained
evidence of livestock. Research conducted on methods of conflict resolution showed that placing
Anatolian Shepherd livestock guarding dogs proved to be effective, with 76% of farmers
reporting a large decline in livestock losses since acquiring an Anatolian. Such solutions appear
effective in increasing farmer’s tolerance for cheetahs, and by the end of the study period cheetah
removals dropped to a mean of 2.1 cheetahs/farm/year. Implementing strategies such as these
could be significant for reducing human-carnivore conflict in the many other places in which it
occurs.

ii
To Khayam,
who showed me the path and provided the vision

Here with Loaf of Bread

beneath the Bough,

A Flask of Wine, a Book

of

Verse - and Thou

Beside me singing in the

Wilderness –

And Wilderness is

Paradise enow.

In: Rubaiyuat of Omar Khayyam

iii
ACKNOWLEDGEMENTS

Although only one name appears on the cover, this thesis would not exist without all
those who have given so generously of their time and energy over the last several years.
In particular, Amy Dickman, my research assistant, who worked diligently by my
side for three years and who shares my interest in science, finding answers through
analysis, and good red wine! Also thanks to Bruce Brewer for managing and building the
Cheetah Conservation Fund’s (CCF) Research and Education Centre while I worked on
this Thesis, and Richard Jeo for his invaluable guidance, and to my International Science
Advisory Board members who gave me continued encouragement during this process and
have mentored me along the way, in particular Linda Munson and Steve O’Brien. I am
indebted to my major supervisors, David Macdonald and Gus Mills who taught me more
than they will know just from this thesis. I am also very grateful to CCF team of staff
and volunteers, in particular Bonnie Schumann, Don Muroura, Matti Nghkembua, and
Susan Deshaies for their valuable input and continued moral support, and to Jack Imbert
our radio-tracking pilot for all the safe hours of flying. In addition, I’m grateful to
Warren Johnson, Christine Breitenmoser-Wursten, and Ruth Feber for their assistance
and editorial comments, and my friends and colleagues at WildCRU who made Oxford
stimulating. In addition, many thanks to my board of directors of CCF-USA, CCF-
Namibia, and CCF-UK for their supporting my time, both financially and morally, while
working on my Thesis, in particular Vance Martin, Cathryn and Carl Hilker, Bill and Pat
Miller, and Charlie Knowles.
This research was conducted with the generous support of the Namibian farmers
who provided access to cheetahs and allowed their release back into the wild, and who
spent hours answering questions about the Namibian farmlands. In particular, Dr. Arthur
Bagot- Smith for first showing me the problems facing the Namibian cheetah, the
Kruger’s for making me apart of their family, and the Waterberg Conservancy farm
families for being great neighbours with a great conservation vision for the future. I am
thankful to the Namibian Government, particularly the Ministry of Environment and
Tourism, which supported this research. And, to His Excellency, Dr. Sam Nujoma, the
President of Namibia and the International Patron for the Cheetah Conservation Fund, for
his interest learning about the Namibian cheetah
In addition, I’d like to acknowledge the various foundations, organisations and
individuals that have supported the work encompassed in this Thesis, particularly major
donors of the Cheetah Conservation Fund in the United States. Partial funding for this
research was provided by the African Wildlife Foundation, the Angel Fund, the American
Zoo Association Conservation Endowment Fund, the Bay Foundation, the Chase
Foundation, Cheetah Conservation Fund Namibia and USA, Cincinnati Zoo, Columbus
Zoo, Earthwatch Institute, the Mazda Wildlife Fund, Philadelphia Zoo, Total SA, the
Weeden Foundation, the WILD Foundation, White Oak Conservation Center and the
WWF SA Green Trust.
On a more personal note, I would like to thank my parents Marline and Ralph
Bushey for teaching me that the world is an open door and one is only constrained to
venture into the world by one’s own fears. They told me I could do anything I put my
mind to.
L.M. - Sept 2002
iv
Table of Contents
CONTENTS

ASPECTS OF CHEETAH (ACINONYX JUBATUS) BIOLOGY, ECOLOGY
AND CONSERVATION STRATEGIES ON NAMIBIAN FARMLANDS

Abstract………………………………………………………………………… ii

Acknowledgements……………………………………………………………. iii

Contents………………………………………………………………………... iv

List of Tables…………………………………………………………………… vii

List of Figures………………………………………………………………….. x

Foreword………………………………………………………………………... xiv
Chapter 1
General Introduction……………………………………………………. 1
Chapter 2
Study Area……………………………………………………………… 24
Chapter 3
Aspects of the Management of Cheetahs Trapped on Namibian
Farmlands………………………………………………………………. 49
Chapter 4
Morphology, Physical Condition and Growth of Namibian Cheetahs…. 75
Chapter 5
Patterns of Molecular Genetic Variation in Namibian Cheetahs………. 92
Chapter 6
A Serosurvey of Antibodies to Viral Diseases in Wild Namibian
Cheetahs………………………………………………………………... 118
Chapter 7
Lymphosarcoma Associated with Feline Leukaemia Virus Infection in
a Captive Namibian Cheetah…………………………………………… 131
Chapter 8
The Incidence of Dental Abnormalities in Wild-Caught Namibian
Cheetahs………………………………………………………………... 138
Chapter 9
Demography of the Namibian Cheetah………………………………… 153
Chapter 10
Notes on the Diet and Feeding Ecology of the Cheetah on Namibian
Farmlands………………………………………………………………. 178
Chapter 11
Movements and Spatial Organisation of Cheetahs on North-Central
Namibian Farmlands: The Influence of Prey Base, Competition and
Perturbation…………………………………………………………….. 199
Chapter 12
Factors Influencing Perceptions and Tolerance towards Cheetahs on
Namibian Farmlands…………………………………………………… 243
v
Table of Contents
CONTENTS (continued)

Chapter 13
Evaluating the Effectiveness of Using Livestock Guarding Dogs as a
Method of Conflict Resolution…………………………………………. 283
Chapter 14
General Discussion and Conservation Implications……………………. 328

Appendices

Appendix I
Current Status of the Cheetah…………………………………… 345
Appendix II
2000/2001 International Cheetah Studbook: Summary…………... 376
Appendix III
Bush Encroachment and Ungulate Density on Commercial
Farmlands in North-Central Namibia……………………………... 397
Appendix IV
Morphometric Protocols Used while Measuring Cheetahs……… 418
Appendix V
Morphometric Data for Cheetahs Published from Various Studies. 419
Appendix VI
Sample Population of Cheetahs used for Genetic Analyses……… 420
Appendix VII
Allele Frequencies for all Loci and Subpopulations……………… 424
Appendix VIII
LOD Scores for Known Dams and Offspring, Showing
Mismatchings and Probability of Non-Exclusion………………… 431
Appendix IX
Questionnaires Used to Interview Farmers……………………….. 433
Appendix X
Questionnaire Used to Evaluate Performance of Livestock
Guarding Dogs……………………………………………………. 435

vi
List of Figures
List of Tables

2.1 Farm sizes in the survey area……………………………………. 30
2.2 Livestock numbers reported in the survey area………………….. 30
2.3 Numbers of wildlife reported in the survey area………………… 31
2.4 Game densities (animals/km2) and estimated biomass (kg/km2)
by habitat type (sparse, medium, thick) using bush canopy……... 37
2.5 Population estimates for three major game species in Namibia…. 41
2.6 Utilisation of cheetah in Namibia 1980 – 1991………………….. 43
3.1 Description of the physical factors used to age cheetahs in this
study……………………………………………………………... 53
3.2 Breakdown of capture events, necropsies and fates of examined
cheetahs…………………………………………………………... 57
3.3 Number of social groups captured and reported each year………. 58
3.4 Demographic composition of cheetahs captured and examined…. 61
3.5 Demographic breakdown of cheetahs examined, separated by
farm type…………………………………………………………. 62
3.6 Frequency of physical or behavioural problems observed in
cheetahs captured and examined………………………………… 63
4.1 The numbers of male and female cheetahs examined, categorised
by age group, during the study period…………………………… 80
4.2 Morphometric data for the wild adult cheetahs (aged over 30 m
at capture, and held in captivity for <30 days), collected using
the protocol shown in Appendix IV……………………………… 82
5.1 Heterozygosity and number of alleles by district………………... 104
5.2 Population pairwise F and R estimates using the combined
st st
data from the regions…………………………………………….. 108
5.3 Relatedness value used to identify possible sire/dam and sibling
relationships within research population of cheetahs……………. 111
6.1 Prevalence of antibodies to selected feline and canine viruses in
free-ranging Namibian cheetahs sampled between 1992-1998….. 125
8.1 Sample population of cheetahs examined for dental
abnormalities……………………………………………………... 144
8.2 Focal palatine erosion scores assigned to captive and wild
cheetahs examined of different ages…………………………….. 144
8.3 Overall frequency of missing premolars, crowded incisors and
perforated FPE in the cheetahs examined………………………... 145
9.1 Annual demographic breakdown into social groups of the
cheetahs captured and examined………………………………… 159
9.2 Information gathered from radio-collared dams regarding
observed reproductive rates, interbirth intervals, litter sizes and
cub mortality……………………………………………………... 166
9.3 Life table showing mortality rates for wild cheetahs throughout
the study period………………………………………………….. 167
10.1 Results of feeding trials performed on captive Namibian
cheetahs………………………………………………………….. 185
vii
List of Figures
10.2 Summary of results from the feeding trials for each prey species
presented………………………………………………………… 186
10.3 Ratios of prey animals consumed, using the corrected scat
analysis, for 100 scats containing prey species………………….. 187
10.4 Contents of wild cheetah scats collected from various locations
on the Namibian farmlands………………………………………. 188
10.5 Ratios of prey animals consumed, calculated using the corrected
scat analysis……………………………………………………… 189
11.1 Mean age, time tracked, number of fixes, minimum distance
moved between fixes, number of fixes to reach first asymptote,
and statistical analyses for radio-tracked cheetahs………………. 213
11.2 Radio-collared cheetahs used in lifetime (95% kernel) home
range analysis 1993 – 2000………………………………………. 214
11.3 Radio collared cheetahs used in annual (95% kernel) home range
analysis 1993 – 2000…………………………………………….. 215
11.4a 95% kernel lifetime yearly home range for different social
groups 1993 – 2000……………………………………………… 218
11.4b Mean home range (lifetime 95% kernel) and statistical analysis
for social groups and sex by seasons - Seasons 1, 2 and 3 are
compared to the dry and wet seasons……………………………. 218
11.5 Minimum density of cheetahs annually – includes all males in
coalition group, and cumulative minimum density of cheetahs
per 1000km2 using all cheetahs tagged and released in study area 225
11.6 Mean habitat preference by social group and sex………………... 228
11.7 Game densities (animals/km2) and estimated biomass (kg/km2)
by habitat type (sparse, medium, thick) using bush canopy……... 229
11.8 Density of cheetahs in this study compared to other
studies…….……………………………………………………… 230
11.9 Data on individual leopards and caracals radio-tracked during
this study…………………………………………………………. 232
11.10 Mean lifetime home range estimations (95% MCP and 95%
kernel for male and female leopards and male caracals.
Statistical differences in lifetime home range estimations are
shown between 95% MCP and 95% kernel home ranges as well
as between sexes and between the wet and dry seasons…………. 233
12.1 Method of calculating the Perception and Tolerance indices……. 251
12.2a Attitudes and removals reported during the baseline 1991-1993
survey…………………………………………………………….. 252
12.2b Attitudes and removals reported during the 1993-1999 survey….. 253
12.2c Statistical analyses of the attitudes and removals between the
1991-1993 and 1993-1999 surveys………………………………. 254
12.3 Comments made by Namibian farmers during the interviews…... 254
12.4a Farmland characteristics reported during the 1991-1993 survey... 256
12.4b Farmland characteristics reported during the 1993-1999 survey... 259
12.4c Statistical analyses of the farmland characteristics between the
1991-1993 and 1993-1999 surveys………………………………. 260
viii
List of Figures
12.5a 1991-1993 survey stocking rates and livestock management
techniques: relationships with cheetah problems and removals…. 264
12.5b Statistical analyses of the stocking rates and livestock
management techniques………………………………………….. 265
12.6a Livestock losses reported during the 1991-1993 survey………… 268
12.6b Livestock losses reported during the 1993-1999 survey………… 269
12.6c Statistical analyses of livestock losses between the 1991-1993
and 1993-1999 surveys…………………………………………... 270
13.1 The sample population of Namibian livestock guarding dogs that
the 334 questionnaires were conducted on during the study,
separated by farm type, age group and sex………………………. 291
13.2 Relationships between the attentiveness, trustworthiness,
protectiveness, farmer satisfaction and care given for the
livestock guarding dogs studied on Namibian farms…………….. 294
13.3 Mean scores for attentiveness, trustworthiness, protectiveness,
care given and farmer satisfaction, for livestock guarding dogs of
different ages on commercial and communal farms……………... 299
13.4 Mean scores for attentiveness, trustworthiness, protectiveness,
care given and farmer satisfaction for male and female livestock
guarding dogs on Namibian farms……………………………….. 300
13.5 Life table for livestock guarding dogs placed on Namibian farms,
using one-yearly intervals………………………………………... 312
A1.1 History of the captive cheetah population……………………….. 353
A1.2 History of the South African captive cheetah population………... 354
A2.1 2000 - 2001 regional groupings of captive cheetahs…………….. 379
A2.2 2000 births by facility……………………………………………. 383
A2.3 2001 births by facility……………………………………………. 389
A2.4 Status of regional populations…………………………………… 396
A3.1 Description of vegetation class by canopy cover………………... 403
A3.2 Comparison of different strip width estimators for large and
small ungulates observed in the study area………………………. 405
A3.3 Results from 2-way analysis of variance regarding the seasonal
habitat preferences of ungulate species………………………….. 407
ix
List of Figures
List of Figures

1.1 Distribution of cheetahs throughout Africa and Asia in 1900, 1975,
and present…………………………………………………………… 10
1.2 Density distribution of Namibian cheetah population……………….. 13
2.1 Land use map of Namibia, showing National Parks, commercial
farms and communal lands…………………………………………... 25
2.2 Namibia rainfall distribution………………………………………… 26
2.3 Vegetation types of Namibia………………………………………… 28
2.4 Map of Namibia, with districts in the North Central Farmlands
highlighted where studies for this thesis were conducted.…………… 29
2.5 The Waterberg Conservancy, an area within the north-central
farmlands, is the centre of the radio-tracking study area…………….. 32
2.6a Annual rain in Waterberg Conservancy cheetah research study area
(1957 – 2000)………………………………………………………... 34
2.6b Mean rainfall per month (averaged from 1957 - 2000) in Waterberg
Conservancy research study area……………………………………. 34
2.7a-b Numbers of individuals of seven game species counted 1995-2000... 36
3.1 Design of a typical capture cage used to live-trap cheetahs on the
Namibian farmlands…………………………………………………. 52
3.2 Number of days between reported date of capture and date of
examination………………………………………………………….. 57
3.3 Reported month of capture for all cheetahs examined………………. 59
3.4 Reported reasons for capture of cheetahs examined………………… 60
3.5 Reported reasons for wild cheetahs entering captivity after
examination………………………………………………………….. 64
3.6 Trends in reported captures due to perceived threats towards game
and/or livestock……………………………………………………… 65
3.7 Reported causes of death for wild cheetahs…………………………. 66
4.1 Ratios of weight to length for adult cheetahs in excellent, fair and
poor physical conditions……………………………………………... 84
4.2 Mean body mass of Namibian cheetahs examined in different age
groups, separated by sex……………………………………………... 85
4.3 Growth curves for a) body mass and b) body length of wild
Namibian cheetah cubs, separated by sex…………………………… 86
5.1 Map of regions where cheetahs originated…………………………... 99
5.2a-b Distribution of alleles (a) and allele frequencies (b) of selected
microsatellites in the 7 regional populations………………………… 105
5.3 Phylogenetic depiction of relationships among 7 subpopulations in
Namibia, constructed using proportion of shared alleles distances
and the neighbour-joining algorithm………………………………… 105
5.4 Population cluster graph from principal component analysis,
showing subpopulation structures……………………………………. 106
5.5 Allele distribution of representative microsatellite loci in the 7
regional groups and the Serengeti population………………………... 107

x