Aspects of Cheetah (Acinonyx jubatus) Biology, Ecology and

Aspects of Cheetah (Acinonyx jubatus) Biology, Ecology and

Aspects of Cheetah (Acinonyx jubatus) Biology, Ecology and

516 Pages ·2003·4.8 MB ·English

Aspects of Cheetah (Acinonyx jubatus) Biology, Ecology and


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



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


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