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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


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