The Black-Footed Cat Felis Nigripes (Burchell, 1824): a Review of the Geographical Distribution and Conservation Status

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THE BLACK-FOOTED CAT FELIS NIGRIPES (BURCHELL, 1824): A REVIEW OF THE GEOGRAPHICAL DISTRIBUTION AND CONSERVATION STATUS

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The user has requested enhancement of the downloaded file. THE BLACK-FOOTED CAT FELIS NIGRIPES (BURCHELL, 1824):

A REVIEW OF THE GEOGRAPHICAL DISTRIBUTION AND

CONSERVATION STATUS

BERYL WILSON

Submitted in partial fulfilment of the requirements for the degree

MAGISTER TECHNOLOGIAE: NATURE CONSERVATION

in the

Department of Nature Conservation

FACULTY OF SCIENCE

TSHWANE UNIVERSITY OF TECHNOLOGY

Supervisor: Dr EP de Crom Co-Supervisors: Prof BK Reilly Dr A Sliwa

May 2015

DECLARATION BY CANDIDATE

"I hereby declare that the dissertation submitted for the degree M Tech: Nature

Conservation, at the Tshwane University of Technology is my own original work and has not previously been submitted to any other institution of higher education.

I further declare that all sources cited or quoted are indicated and acknowledged by means of a comprehensive list of references."

BY Wilson

This study is dedicated to my late father

Ron Wilson

who had no idea why or how this study was done,

but nevertheless supported me 110% in his own way.

May I be at least half as successful as he thought I was!

iii

ACKNOWLEDGEMENTS

I would like to express my sincere gratitude and appreciation to the following persons or institutions, in no particular order:

 My supervisors, Dr Nellie de Crom and Prof Brian Reilly, for stepping in so ably

at the last moment and picking up the many pieces and promises left behind by

others before them.

 Department Sports, Arts and Culture: Northern Cape for the initial registration

funding.

 The McGregor Museum and staff for coping with the poor timing of my final

write-up and my absence during a trying transitional period for the institute.

 Black-footed Cat Working Group members for their support, motivation,

inspiration and for being there through some incredible experiences in the field.

 Dr Alex Sliwa for the initial concept of a distributional study and for developing

the project’s public awareness posters, as well as his support and advice

throughout.

 Marion Holmes, founding trustee of the Cat Conservation Trust, for her

investigations into taxidermy records in the early years of the investigation.

 Citizen scientists, taxidermists and predator control professionals for sharing

localities and sightings, many confidentially and without whom this study would

not have been possible.

 Dr Lizanne Roxburgh, senior scientist from the Endangered Wildlife Trust for

leaping into action at the very last minute to assist in producing the final

population density map when my GIS skills faltered.

 Luger Wilson, my constant companion, assistant and watchdog throughout.

ABSTRACT

The Black-footed Cat (Felis nigripes) is one of the rarest cat species in Africa and is restricted to the southern African subregion. A review of historical and current geographical distribution ranges was made from 2006 to 2014. Locality records

(790) were assembled from literature reviews, museum records, field surveys, web-based resources and citizen scientist observations. Maps were digitised and analysed using Geographical Information Systems. Historical geographical distributions contained a limited number of records and underestimated the range of the species. The current mapping suggested a wider but more fragmented geographical range than previously assumed. A replicable mapping method was developed to estimate population sizes in South Africa and regionally to allow for more informed decision-making in future conservation assessments. Until recently, the species had a South African national conservation status of Least Concern that is in the process of being uplifted to Vulnerable. The findings here support this change in conservation status, as well as the continued IUCN global listing of

Vulnerable. Apart from the already acknowledged threats to the species, an additional eleven conservation risks were identified and ranked according to severity. This information was used to make management recommendations and identify research priorities for the species.

CONTENTS

PAGE

DECLARATION BY CANDIDATE ii

DEDICATION iii

ACKNOWLEDGEMENTS iv

ABSTRACT v

LIST OF FIGURES xii

LIST OF TABLES xvii

GLOSSARY xviii

CHAPTER 1

INTRODUCTION AND AIMS OF THE STUDY 1

1.1 INTRODUCTION 1

1.2 THE STUDY REGION 3

1.3 RATIONALE 6

1.4 RESEARCH DEVELOPMENT BACKGROUND 9

1.5 OBJECTIVES AND KEY QUESTIONS 9

1.5.1 Geographical distribution: To determine more accurately than

previously the geographical distribution of the black-footed cat 10

1.5.2 Conservation status: To determine the current conservation of

black-footed cat 10

1.6 METHODOLOGY OUTLINE 11

CHAPTER 2

SPECIES REVIEW 12

2.1 INTRODUCTION 12

2.2 TAXONOMY 12

2.3 DESCRIPTION 14

2.4 BEHAVIOUR 17

2.5 DIET 18

2.6 BIOLOGY 19

2.7 GEOGRAPHICAL DISTRIBUTION 21

2.8 CONSERVATION STATUS 22

2.9 PRINCIPAL THREATS 23

CHAPTER 3

RESEARCH METHODS 24

3.1 INTRODUCTION 24

3.2 RESEARCH DESIGN 24

3.3 DATA COLLECTION 25

3.3.1 Literature review 25

3.3.2 Museum records 26

3.3.3 Web-based resources 27

3.3.4 Observations 28

3.3.4.1 Field surveys 28

3.3.4.2 Citizen scientist records 29

3.4 DATA ANALYSIS 32

3.4.1 Geographical records 32

vii

3.4.2 Mapping 35

3.4.3 Statistical method 36

3.4.4 Population size estimates 36

3.5 LIMITATIONS 37

3.5.1 Observer biases 38

3.5.2 Historical imprecision 40

3.5.3 Record verification 41

3.5.4 Gaps in data 41

3.6 ETHICAL CONSIDERATIONS 42

3.7 DISSERTATION OVERVIEW 43

CHAPTER 4

RESULTS AND DISCUSSION: GEOGRAPHICAL DISTRIBUTION 44

4.1 INTRODUCTION 44

4.2 REVIEW OF SURVEY RESULTS 45

4.2.1 Record and observer categories 45

4.2.1.1 Sight records 46

4.2.1.2 Specimen records 47

4.2.1.3 Captive records 48

4.2.1.4 Capture-release records 48

4.2.1.5 Roadkill records 49

4.2.1.6 Fossil records 50

4.2.2 Spatial distribution of records 51

4.2.3 Temporal distribution of records 53

4.3 REVIEW OF HISTORICAL MAPS 55

viii

4.4 CURRENT MAPPING 60

4.4.1 Observer rating (OR) maps 61

4.4.2 Historical records (up until the end of 1989) 64

4.4.3 Current records (1990 onwards) 67

4.4.4 Current geographical distribution of black-footed cat 69

4.5 DISCUSSION 71

4.5.1 Limitations of data and some assumption 71

4.5.1.1 Observer effects 71

4.5.1.2 Data gaps 72

4.5.1.3 Research effort 74

4.5.2 Range extensions 75

4.6 SUMMARY 80

CHAPTER 5

RESULTS AND DISCUSSIONS: CONSERVATION STATUS 81

5.1 INTRODUCTION 81

5.2 ESTIMATION OF POPULATION SIZE 82

5.3 CURRENT CONSERVATION STATUS 85

5.3.1 Global and regional status 85

5.3.2 National status 86

5.3.3 Principal conservation threats 87

5.4 EMERGING ISSUES 88

5.4.1 Road mortalities 88

5.4.2 Hybridisation 90

5.4.3 Population genetics 90

5.4.4 Fragmented distribution 91

5.4.5 Habitat preferences 92

5.4.6 Indiscriminate persecution 93

5.4.7 Industry interest and use 94

5.4.8 Reliance on sympatric species 97

5.4.9 Intraguild predation 99

5.4.10 Natural catastrophes 102

5.4.11 Diseases 104

5.4.12 Climate change 105

5.4.13 Presence in formally protected areas 106

5.5 DISCUSSION 109

5.5.1 Range and population size 109

5.5.2 Anthropogenic threats 112

5.5.3 Natural threats 118

5.5.4 Ranking of the threats 128

5.6 SUMMARY 130

CHAPTER 6

CONCLUSIONS AND RECOMMENDATIONS 132

6.1 INTRODUCTION 132

6.2 SYNTHESIS OF THE FINDINGS 132

6.2.1 Geographical distribution 132

6.2.2 Conservation status 134

6.3 CONSERVATION IMPLICATIONS 135

6.4 MANAGEMENT RECOMMENDATIONS 138

6.5 FUTURE RESEARCH PRIORITIES 140

REFERENCES 143

LIST OF ANNEXURES

ANNEXURE A: PUBLIC AWARENESS POSTER: ENGLISH 166

ANNEXURE B: PUBLIC AWARENESS POSTER: AFRIKAANS 167

ANNEXURE C: QUESTIONNAIRE: COMPLETED EXAMPLE 168

ANNEXURE D: ETHICAL CLEARANCE 169

LIST OF FIGURES

PAGE

FIGURE 1.1: A male black-footed cat (Felis nigripes), research subject

“Pogo” 1

FIGURE 1.2: The study area in southern Africa (QGIS, 2015) 4

FIGURE 1.3: Country-based species richness of small carnivores

(excluding Felidae and Canidae) in Africa (Do Linh San et

al., 2013:8) 6

FIGURE 1.4: A typical observation of the naturally rare, small sized,

highly elusive and cryptic black-footed cat is often limited to

a brief sighting only 7

FIGURE 2.1: A black-footed cat showing the typical patterning and

colouration, as well as the short tail 15

FIGURE 2.2: A leaping black-footed cat showing the black undersides of

its feet 16

FIGURE 2.3: Captive-bred black-footed cat kittens at the Cat

Conservation Trust playing in a hollowed-out Agave stump

similar to a natural termitarium 19

FIGURE 4.1: Black-footed cat survey results grouped according to type

of record and observer rating categories 46

FIGURE 4.2: A black-footed cat unintentionally trapped by a farmer in a

cage trap set for larger problem-causing predators which

was later released after being correctly identified 49

xii

FIGURE 4.3: A sequence of photographs showing a male black-footed

cat (BFCWG research subject “Kubu”) deliberating and

successfully negotiating busy traffic on a national road (N8)

near Kimberley, Northern Cape, South h Africa

(Photograph: A. Sliwa) 50

FIGURE 4.4: A lateral view of a black-footed cat jaw fragment (UW 88-

517) recovered from Malapa Hominin site, South Africa

(Kuhn et al., 2011:7) 50

FIGURE 4.5: Spatial distribution of black-footed cat survey results

grouped by country 51

FIGURE 4.6: Spatial distribution of black-footed cat survey results

grouped by South African province 53

FIGURE 4.7: Temporal distribution of black-footed cat distribution

records 54

FIGURE 4.8: Black-footed cat distribution (approximately 1 236 178 km2)

as presented in Stuart (1982:8) 56

FIGURE 4.9: Black-footed cat geographical distribution (approximately

1 253 534 km2) as presented in Smithers (1983:392) 56

FIGURE 4.10: Black-footed cat geographical distribution (approximately

1 401 004 km2) as presented in Skinner and Smithers

(1990:419) 57

FIGURE 4.11: Black-footed cat geographical distribution (approximately 1

530 437 km2) as presented in Skinner and Chimimba

(2005:407) 57

xiii

FIGURE 4.12: Black-footed cat geographical distribution (approximately

2 043 713 km2) according to Wilson and Sliwa (2007) 58

FIGURE 4.13: Changes in the area size (km2) of the envisaged black-

footed cat geographical distribution as described in

publications from 1982 to 2007 59

FIGURE 4.14: Black-footed cat records with an unreliable record rating

(OR 1) indicated as red points 61

FIGURE 4.15: Black-footed cat records with a single, reliable rating (OR

2) indicated as blue points 62

FIGURE 4.16: Black-footed cat records with a multiple, reliable rating (OR

3) indicated as black points 63

FIGURE 4.17: Combined black-footed cat observer rating distribution

records (with unreliable OR 1 indicated as red points;

single but reliable OR 2 indicated as blue points, and

multiple and reliable OR 3 indicated as black points) 64

FIGURE 4.18: Historical (up until the end of 1989) black-footed cat

distribution records indicated as purple triangles 65

FIGURE 4.19: Composite map of black-footed cat historical (up until the

end of 1989) geographical distribution (Stuart (1982) in

pink; Smithers (1983) in green, and Skinner and Smithers

(1990) in purple 66

FIGURE 4.20: A composite map of black-footed cat historical (up until the

end of 1989) geographical distribution with Skinner and

Chimimba (2005) in deep yellow, and Wilson and Sliwa

(2007) in pale yellow 67

xiv

FIGURE 4.21: Current (from 1990 onwards) black-footed cat distribution

records indicated as green diamonds 68

FIGURE 4.22: Combined black-footed cat observer rating distribution

records (with unreliable OR 1 indicated as red points;

single but reliable OR 2 indicated as blue points, and

multiple and reliable OR 3 indicated as black points) with a

proposed current geographical distribution outline overlaid

as dashed green line 69

FIGURE 4.23: Current proposed basic black-footed cat geographical

distribution map (approximately 2 214 267 km2) 70

FIGURE 5.1: A kernel density estimate heatmap of black-footed cat

distribution patterns in southern Africa (Map: L. Roxburgh) 83

FIGURE 5.2: A black-footed cat roadkill victim near Aberdeen submitted

as a sighting record by a citizen scientist 89

FIGURE 5.3: A black-footed cat on roadside display in the Karoo after

being trapped and killed after raiding a chicken coup 94

FIGURE 5.4: The researcher hosts scientific safaris for numerous

groups and local television news crews in a bid to raise

awareness about the black-footed cat 97

FIGURE 5.5: A black-footed cat kitten in a hollowed out snouted

harvester termite (Trinervitermes trinervoides) mound

(Photograph: A. Sliwa) 98

FIGURE 5.6: Black-footed cat geographical distribution (blue) overlying

the range of springhare (grey) indicating am 87.6%

intersection with black-footed cat range 99

FIGURE 5.7: The discarded and uneaten carcass of “Maya” a young

female black-footed cat killed by a caracal in 2007 100

FIGURE 5.8: “Maya”, a young female black-footed cat killed by a caracal

in 2007 showing bite wounds circled in red 101

FIGURE 5.9: The only remains of “Panga”, a large radio-collared male

black-footed cat that was killed and eaten by black-backed

jackals in 2007 101

FIGURE 5.10: The remains of “Gogo”, a radio-collared female black-

footed cat that died aboveground, possible due to a severe

storm in 2009 103

FIGURE 5.11: The remains of “Jimbo”, a radio-collared male black-footed

cat that died in a den that collapsed after it was flooded in

2009 (Photograph: A. Sliwa) 104

xvi

LIST OF TABLES

PAGE

TABLE 5.1: Estimated areas of high, medium and low density black-

footed cat regions, and population size both nationally and

globally 84

TABLE 5.2: Formally protected areas in southern Africa indicating the

presence/absence of black-footed cat within the boundaries,

or within a 50 km radius of the nearest black-footed cat

locality record 107

TABLE 5.3: A summary of the conservation threats to black-footed cats

(Felis nigripes) ranked in order of decreasing severity 129

TABLE 6.1: A summary of proposed conservation interventions for the

black-footed cat (Felis nigripes) ranked in order of priority 139

xvii

GLOSSARY

AOO Area of occupancy: the area inside of the EOO were a

species actually occurs

BFC Black-footed cat/s (Felis nigripes) (Burchell, 1824)

BFCWG Black-footed Cat Working Group

EOO Extent of occurrence: the minimum convex polygon

encompassing all known normal occurrences of a particular

species

Extant A species that is still in existence, i.e. not extinct

CITES The Convention on International Trade in Endangered

Species of Wild Fauna and Flora, an international

agreement between governments that aims to ensure that

the international trade in specimens of wild animals and

plants does not threaten their survival

FPA Formally protected area: a national park or nature reserve

that is managed as a proclaimed wildlife preserve by a

national or provincial authority

Geographical The natural arrangement of a species in a region, area or distribution specific locality

Grey literature Technical reports from provincial government research

reports, working papers from research groups, etc. which

have not been published commercially or widely accessible

but which are still important sources of information

xviii

IUCN International Union for Conservation of Nature, the world’s

oldest and largest global environmental organisation

Inquilinism A commensal situation in which one species lives in the

nest, burrow, or dwelling place of another animal without

harming the host

Intraguild The killing and eating of species that use similar resources predation are thus potential competitors

KDE Kernel density estimation - a non-parametric way to

estimate the probability density function of a random

variable

Obligate carnivore Animals that depend solely on animal flesh for their nutrient

requirements

QDS Quarter Degree Square – a way of dividing longitude-

latitude degree square cells into smaller squares of 15 x 15

minutes and assigning a unique geocode e.g. 2824 Db and

typically representing an area of 27.4 km x 25km (or

685km2) in southern Africa

Range The maximum area that a species is expected to potentially

occupy

xix

CHAPTER 1

INTRODUCTION AND AIMS OF THE STUDY

1.1 INTRODUCTION

The black-footed cat (Felis nigripes) (Burchell, 1824) (Figure 1.1), with a highly restricted range in southern Africa, is a vulnerable species and one of Africa’s three endemic feline species (Macdonald, Loveridge & Nowell, 2010:5). According to the IUCN (the International Union for Conservation of Nature), as of 2015, there are considered to be 38 extant feline species globally. Of these species, 16 are included in the top three categories of the IUCN Red List of Threatened Species

(Macdonald, Loveridge & Nowell, 2010:15). Further to this, of the ten African species, two are considered vulnerable and three are near threatened.

FIGURE 1.1: A male black-footed cat (Felis nigripes), research subject “Pogo”

The majority of threats facing carnivores are anthropogenic in nature, with the leading threat being habitat loss due the world’s rapid human population growth and resulting conversion of natural vegetation into cultivated areas (Loveridge,

Wang, Frank & Seidensticker, 2010:162). Add to this, non-natural mortalities from poaching, illegal hunting and trapping, vehicle collisions, problem animal control

(legal and illegal) and disease exposure (Loveridge et al., 2010:164) and the situation is dire. Fortunately, there is a strong commitment to carnivore conservation in most developed countries (Loveridge et al., 2010:178), and it is becoming increasingly recognised that the success of conservation efforts are inextricably linked to the social, political and cultural landscapes in which intervention measures are implemented (Akerlof & Kennedy, 2013:11).

Whilst a few of the larger cats are familiar to most people, the large majority of cat species of the world remain unknown to the average person which is also reflected in the amount of research effort on small carnivores in Africa (Do Linh San &

Somers, 2013:1). According to their findings, carnivore research effort is largely driven by body size and range size. With black-footed cats (referred to as BFC throughout this dissertation) being the smallest of the African species (Sliwa,

Herbst & Mills, 2010:538) and having the most restricted range (Nowell & Jackson,

1996:8), the paucity of information on this species is understandable.

According to Sanderson, founder and director of the Small Wild Cat Conservation

Fund1 in Shea (2014), this imbalance extends to funding too. Since 2007, the seven iconic big cat species received 99.22% of all funding for cat conservation,

1 www.smallcats.org 2

with the known 22 small cat species around the world only receiving a paltry

0.78%.

Throughout the literature reviewed, the BFC has been described as uncommon to rare (Shortridge, 1934:96; Smithers, 1971:128; Visser, 1976:73; Stuart, 1982:8;

Stuart & Wilson, 1988:23; Nowell & Jackson, 1996:8; Sunquist & Sunquist,

2002:80; Skinner & Chimimba, 2005:406; Sliwa, 2013:204). This was the basis for most researchers explaining the general lack of distributional data available for the species, as well as the subsequent differing estimations of geographical range.

However, knowledge of geographical distributions, habitat preferences and population sizes are central to the conservation and management of any species.

The rise of new powerful statistical techniques and Geographical Information

System (GIS) tools has enhanced and improved the accuracy of range predictions of organisms (Guisan & Zimmerman, 2000:148). Pearson, Raxworthy, Nakamura and Peterson (2007:107) demonstrated the success of predicting species distributions from small numbers of occurrence in records using cryptic geckos in

Madagascar as a test case. By employing these new tools and techniques, it may be possible to establish a more current and accurate geographical distribution for the BFC using a limited number of citizen science and historical records.

1.2 THE STUDY REGION

The study was restricted to the southern African subregion of the African continent as this is the known range of the species (Stuart, 1982:8; Stuart & Stuart,

1988:152; Smithers, 1983:392; Skinner & Smithers, 1990:419; Friedman & Daly,

2004:173; Skinner & Chimimba, 2005:407; Wilson & Sliwa, 2007). The geographical subregion consists of numerous countries including South Africa,

Namibia, Botswana, Lesotho and Swaziland, but also encompassed the surrounding territories of Zimbabwe, Mozambique, Angola and Zambia (Figure

1.2) from -12° S of the equator and downwards to the southern tip of Africa for the purposes of this study.

FIGURE 1.2: The study area in southern Africa (QGIS, 2015)

The southern African subregion has a wide diversity of ecoregions and can be broadly grouped into the following dominant biomes namely grasslands, woodlands, shrubland, savanna and desert (UNEP, 2008:10).

Much of the landscape has been transformed to meet the social and agricultural needs of the rapidly growing human populations with sub-Saharan Africa that are expected to continue to add to the world population up until 2050 than any other region (Haub & Kaneda, 2013:6). However, droughts are common in the western areas and these impact significantly on the agroecological potential of much of the subregion.

Much of the subregion’s climate is influenced by the warm Mozambique currents in the Indian Ocean and the cold Benguela currents in the Atlantic Ocean. Rainfall over the southern African subregion varies from over 1 000 mm in the east to less than 200 mm in the west. Floods and prolonged droughts are typical climatic conditions (UNEP, 2002:32). Temperatures in the subregion differ dramatically across the range and locally. Overall, the eastern regions tend to experience less severe mean temperature fluctuations between seasons, whilst the western areas have low winter and very high summer temperatures (UNEP, 2008:8).

South Africa, home to the largest proportion of the geographical distribution of the

BFC, is the third most biologically diverse country in the world after Indonesia and

Brazil (Wynberg, 2002:233-243). According to Do Linh San and Somers (2013:2),

Africa’s small carnivores represent about 34% of extant small carnivores, with the southern African subregion represented by at least 11-15 species (Figure 1.3).

However, it should be noted that their assessment did not include species from the

Canidae and Felidae groups as these assessments are covered by other IUCN

Species Specialist Groups (Schipper, Hoffmann, Duckworth & Conroy, 2008:29).

However, since BFC are also tiny carnivores, it could be argued that they share

many of the challenges and threats facing the Mustelidae, Nandiniidae,

Herpestidae and Viverridae groups.

FIGURE 2.3: Country-based species richness of small carnivores (excluding Felidae and Canidae) in Africa (Do Linh San et al., 2013:8)

1.3 RATIONALE

The historical paucity of data on this species has led to inconsistencies and perpetuated inaccuracies in current literature that in turn may be affecting the

accuracy of conservation measures. Its unique attributes and characteristics have contributed to these inaccuracies (Figure 1.4).

FIGURE 1.4: A typical observation of the naturally rare, small sized, highly elusive and cryptic black-footed cat is often limited to a brief sighting only

However, this is not unexpected as research efforts on smaller carnivore species are often reduced by intrinsic and extrinsic factors (Brodie, 2009:2927; Brook,

Bielby, Nambiar & Carbone, 2014:2). In Africa, some of the smaller, diurnal, social carnivore species have received some attention, but for the most part, the behaviour, ecology and ecological roles of most African small carnivores remains unknown (Do Linh San et al., 2013:5). Nowell and Jackson (1996:196) specifically stated that for many of the most vulnerable small cats, there is little knowledge of their biology and ecology making conservation of these species difficult. According to Brooke et al. (2014:5), the mean number of papers per Felidae species 7

published between 1900 and 2010 is 74.20. All African Felis species are noted by them as having received little research effort.

The literature research for this review suggested that, in particular for BFC, research effort was very low i.e. less than 30 publications on the species in the same period (1900-2010) with the majority published from 1960 onwards.

Furthermore, only seven research papers or references in the past 35 years specifically addressing the full geographical range of BFC, many of which contained only basic range maps (Stuart, 1982:8; Smithers, 1983:392; Stuart &

Stuart, 1988:152; Skinner & Smithers, 1990:419; Friedman & Daly, 2004:173;

Skinner & Chimimba, 2005:407; Wilson & Sliwa, 2007). These geographical distributions are usually measured as the extent of occurrence (EOO) which is thought of as the minimum convex polygon encompassing all known normal occurrences of a particular species and is the range usually depicted in field guides. However, a species cannot be expected to occupy the entire EOO as it is likely to contain unsuitable or unoccupied habitat. Therefore, EOO is emphatically not a measure of the area over which a species is actually found to occur (Gaston

& Fuller, 2009:4).

The process of ranking species vulnerability by the International Union for

Conservation of Nature (IUCN)2 uses a number of criteria with the size of the remaining population and its geographic range being the main underlying factors that influence population size and extinction risk (Nowell & Jackson, 1996:2-6). For

2 www.iucn.org/ 8

this reason, it is essential that the most accurate range baseline data is employed from the outset.

1.4 RESEARCH DEVELOPMENT BACKGROUND

This study was initiated by the Black-footed Cat Working Group (BFCWG) comprising of a number of principal investigators in various disciplines. Following the first ecological study on the species on Benfontein Nature Reserve, near

Kimberley, South Africa by Sliwa from 1992-1998, the group was established in

2007 to broaden the study. Members of the group have since conducted various field-based investigations including general ecology, reproductive biology, geographical range, habitat preferences, health and conservation of the species.

The preliminarily results of these studies that bear reference to this study have been included later in this research.

This study, conducted from 2006-2014 (but ongoing), focuses on determining the current geographical distribution of the BFC, and attempts to understand how this has changed from any known historical data. The implication of changes (if any) may provide an empirical basis for future conservation assessments to be based on.

1.5 OBJECTIVES AND KEY QUESTIONS

Broadly speaking, the objective of this study focussed on improving the accuracy of the historical and the current geographical distributions of the BFC. This

information can be used to predict future changes and to identify additional conservation threats to the species that in turn can lead to more appropriate future assessments of the species. The two main objectives and the respective key questions are as follows:

1.5.1 Geographical distribution: To determine more accurately than

previously the geographical distribution of the black-footed cat.

Key questions:

 How accurate were the previous extent of occurrence geographical distribution

estimations for the black-footed cat?

 What is the current extent of occurrence of the black-footed cat?

 How (if at all) does the current geographical distribution of the black-footed cat

compare to the previously documented historical ranges?

1.5.2 Conservation status: To determine the current conservation status of

the black-footed cat.

Key questions:

 Is it possible to estimate a population size for the black-foot cat using the

current geographical distribution information?

 In what ways will the new geographical information and mapping method assist

with conservation protection measures for the species?

 What are the additional conservation risks suggested by the new information

about the geographical distribution of the black-footed cat?

1.6 METHODOLOGY OUTLINE

Over 800 records were obtained during the study period, but data collection is ongoing under the auspices of current BFCWG activities. Records were a combination of historical and current records obtained from personal observations,

BFCWG field activities, books, travel accounts, museum collections, published reports, journal, books, interviews, public sightings, citizen scientist databases, internet online forums, conservation surveys, taxidermists, and SAPS case files.

The data was then checked for reliability and rated accordingly and then plotted into various mapping systems and formatted for GIS analysis and mapping.

Limitations considered were, but not limited to, observer biases, historical imprecision, record verification and various spatial and temporal gaps in the data.

CHAPTER 2

SPECIES REVIEW

2.1 INTRODUCTION

The black-footed cat (BFC) (Felis nigripes), also known as the small-spotted cat, is an extant, small vulnerable felid restricted to the more arid and semi-arid parts of southern Africa. It is Africa’s only endemic Felis species (Macdonald, Loveridge &

Nowell, 2010:5). There is a paucity of information on all aspects of its biology, particularly accurate data about its current distributional range and habit requirements. This may be one of the more serious threats to future conservation assessments.

2.2 TAXONOMY

First described by Burchell (1824:592), the BFC is part of the Felis group that is now restricted to members of the domestic cat lineage, some of the smallest species of the family (Sliwa, 2013:196). It belongs to the lineage of Old World domestic cats currently divided into four species, F. nigripes, F. silvestris, F. margarita and F. chaus, (Werdelin, Yamaguchi, Johnson & O’Brien, 2010:61) of which the latter two are extralimital to southern Africa (Bronner, Hoffman, Taylor,

Chimimba, Best, et al., 2003:72).

Werdelin et al. (2010:72) stated that there were no positive fossil records for Felis nigripes. This statement is, however, inaccurate because the first fossil records of

the species were described by Klein, Cruz-Uribe and Beaumont (1991:99). These fossils were recovered from Equus Cave near Taung in Northern Cape Province, which is in close proximity to the site of the Burchell type specimen. Radio carbon dating placed the age of these fossils from three periods spanning 18 000 - 6 000

BP (Plug & Badenhorst, 2001:87). More recently, Kuhn, Werdelin, Hartstone-

Rose, Lacruz and Berger (2011:7) recovered a significantly older fossil of about

1.977 million years old from the Malapa Hominin site near Krugersdorp, Gauteng

Province.

Two subspecies of BFC have been described and these persist in theory

(Meester, Rautenbach, Dippenaar & Baker, 1986:130-131) having been neither confirmed nor discounted genetically to date. Felis nigripes nigripes, the nominate genotype form described by Burchell, was first described from near Kuruman in the Northern Cape. Its range includes present-day south-eastern Namibia,

Botswana, and the South African provinces of the Northern Cape, North West,

Gauteng, Limpopo and marginally into Mpumalanga. Felis nigripes thomasi

Shortridge, 1931 (Shortridge, 1931:119) is described from a specimen collected near Grahamstown in the Eastern Cape. Its range includes the present-day

Eastern Cape, westwards to the southern regions of the Northern Cape and the

Free State.

However, Smithers (1971:128) doubted the validity of two subspecies, and this is thought is shared by Sliwa (2013:203) who suggests that the species may be polymorphic in the centre of range but does exhibit some geocline variations towards the extremes of its range. The northern race typically tends to be paler

with less distinct striping whereas the southern race has a tawnier appearance with bolder patterning. However, since there are no oblivious geographical or ecological barriers between the ranges which are normally the leading causes of genetic drift and speciation (Hoskin, Higgie, McDonald & Moritz, 2005:1353), it is likely that the subspecies status is invalid.

Both Shortridge (1934:96) and Rautenbach (1982:150) suggested possible hybridisation with domestic cats. Rautenbach further mentioned an example of a hybrid cat “seen in the business centre of Pretoria on a Saturday morning”.

However, Král and Zima (1980) in Wozencraft (1993:536) noted a distinctly different karotype from other Felis, and Lopez, Culver, Stephens, Johnson and

O’Brien (1997:281) reported distinct DNA sequences of this species that set it apart from the other Felis spp. concluding that the species evolved separately from an early point of the group’s establishment.

It is likely that so-called hybrid reports in literature were mistaken identifications.

Juvenile African wildcats (Felis silvestris lybica) share a remarkable resemblance with BFC until the intense spotting fades and paler, uniform adult coloration develops. The only confirmed hybrid cases (BFC and domestic cats) were specifically forced under captive conditions by Leyhausen (1979:95).

2.3 DESCRIPTION

The BFC is the smallest African cat (Skinner & Smithers, 1990:416; Sliwa, Herbst

& Mills, 2010:538) and the second smallest cat species in the world (Macdonald,

Loveridge & Nowell, 2010:3). Stocky-bodied, males have an average mass of 1.9 kg and females of 1.3 kg (Sliwa, 2004:99). It is boldly patterned with blackish spots

(hence the alternative name, small-spotted cat) on a tawny-cream background

(Figure 2.1).

FIGURE 2.1: A black-footed cat showing the typical patterning and colouration, as well as the short tail

The legs are barred with thick, transverse dark stripes. The undersides of the feet are black (Figure 2.2), a feature shared by the African wildcat. However, unlike wildcats, its legs appear proportionally shorter in comparison to the body. Also distinctive, is the noticeably shorter tail that is <40% of the length of the head and body (Sliwa, 2013:203).

FIGURE 2.2: A leaping black-footed cat showing the black undersides of its feet

2.4 BEHAVIOUR

Adapted to hunting in short vegetation, these cats are predominantly ground- dwellers, and will not readily take to trees like other cat species. They lead a solitary existence except when with kittens or during brief mating periods. The

BFC is strictly crepuscular and nocturnal. They are active throughout the night and will even hunt at temperatures of -8 ˚C (Olbricht & Sliwa, 1997:86). Brief periods may be spent basking in the sun close to their dens during colder periods.

Black-footed cats are easily startled and flee readily but will fight ferociously when cornered or handled. A Khoisan (Bushman) legend tells of a cat that took down a giraffe (Giraffa camelopardalis) by piercing its jugular emphasising the legendary courage contained in such a small cat (Sunquist & Sunquist, 2002:78).

Only one social organisational and spatial system study has been published on the species (Sliwa, 2004:96-100) that concluded with the following information. Home ranges, when incorporating all locations (100 % minimum convex polygon), were stable and averaged 20.7 km2 for males and about 10 km2 for females with some partial overlapping. Since range size is dependent on available prey resources, in more arid regions these home ranges can be considerably larger. Both sexes spray mark, particularly during mating season when they are deployed in proportion to intensity of use and may play a role in social spacing (Molteno, Sliwa

& Richardson, 1998:40).

During the day, BFC make use of dens. The species prefers hollowed out abandoned termite mounds when available (especially for the kittens), but will use dens dug by other animals such as springhares (Pedetes capensis), Cape ground squirrel (Xerus inauris) and aardvark (antbear) (Orycteropus afer) Wilson (pers. obs.).

2.5 DIET

As with most small cats, BFC are obligate carnivores. According to Sunquist and

Sunquist (2002:78), prior to Sliwa’s field study published in 2006, the only information about the diet of BFC came from specimens that had been either shot or were roadkill victims. In this study, Sliwa noted that these cats were capable of catching relatively large prey up to the size of Cape hare (Lepus capensis) and

Northern black korhaan (Afrotis afraoides). However, they were mostly opportunistic feeders with 98% of their diet consisting of small mammals (72%) and birds (26%) and inclusive of more than 55 vertebrate and invertebrate recorded species (Sliwa, 2006:198-199) in the Northern Cape province alone. This diet is likely to be regionally and seasonally dependent. The cats are extremely active in search of food travelling 4.5 - 16 km per night and will kill 10 - 14 items per day representing about 250 - 300 g of food (Olbricht & Sliwa, 1997:85). Prey items are either stalked or ambushed. Cats have also been observed scavenging dead springbok lambs during lambing season (Sliwa, 1994:91). Healthy cats are independent of drinking water (Sliwa, 2013:295).

2.6 BIOLOGY

Black-footed cats mate year-round in captivity but in the wild they exhibit a distinct a season from late winter (August) to March (Olbricht & Sliwa, 1997:84). Up to two litters a year may be produced, with one to four kittens (normally two) born after a

63 - 68 day gestation inside a springhare burrow or hollow termite mound (Olbricht

& Sliwa, 1997:84; Skinner & Chimimba, 2005:408). Births are timed to coincide with rains and food availability. Kittens (Figure 2.3) become independent at around three-four months but will remain within range of their mother for extended periods

(Sliwa, 2013:205).

FIGURE 2.3: Captive-bred black-footed cat kittens at the Cat Conservation Trust playing in a hollowed-out Agave stump similar to a natural termitarium

Whilst they are reported to live up to 16 years in captive situations (Sliwa,

2013:205), life expectancy in the wild is about five years (BFCWG, unpublished data) although two monitored individuals lived for at least seven years (Sliwa,

Wilson, Lamberski, Lawrenz & Herrick, 2011; Sliwa, Wilson, Küsters, Lawrenz,

Eggers, et al., 2015).

Young or sick individuals are vulnerable to intraguild predation by black-backed jackals (Canis mesomelas) and caracals (Caracal caracal), as well as domestic dogs (Canis familiaris) and even Anatolian shepherd dogs (Wilson, pers. obs.).

In captivity, BFC show a high prevalence for AA-amyloidosis (Terio, O’Brien,

Lamberski, Famula & Munson, 2008:393). This is a disease characterised by fibrillar protein depositions in many organs usually cumulating in renal failure.

About 70% of the documented deaths of captive cats internationally are because of this disease, much of it presumably linked to captive conditions (Terio, et al.,

2008:396). However, the presence of amyloid in a free-ranging BFC was detected by Terio et al. (2008:396) and Zimmermann, Lawrenz and Sliwa (2011:364) providing additional evidence for a species predilection and supporting the existence of a possible familial type of amyloidosis in the BFC.

Various ectoparasites (Horak, Heyne & Donkin, 2010:7) and endoparasites (Sliwa,

2013:205) have been recorded, and studies to determine infectious disease prevalence in the cats and other small sympatric carnivores are underway

(Lamberski, Sliwa, Wilson, Herrick & Lawrenz, 2009). Amongst small carnivores sampled from the same locality, namely Benfontein Game Reserve near

Kimberley, the ectoparastitic burden was highest in BFC (Matthee, van der

Mescht, Wilson & Lamberski, 2011:29).

2.7 GEOGRAPHICAL DISTRIBUTION

Black-footed cats are considered to have the most restricted distribution of any of the African species (Nowell & Jackson, 1996:8). The species is endemic to the arid grasslands, dwarf shrub, and savanna of the Karoo and Kalahari in southern

Africa (Sliwa, 2013:204). However, over the past 40 years, various researchers

(Stuart, 1982:8; Smithers, 1983:392; Stuart & Stuart, 1988:152; Skinner &

Smithers, 1990:419; Friedman & Daly, 2004:173; Skinner & Chimimba, 2005:407;

Wilson & Sliwa, 2007) have reported and published substantially different range occurrences. This has complicated the determination of an accurate geographical distribution in the past, which may in turn have detrimentally influenced published field guides and conservation assessments to date.

Furthermore, the predicted geographical distributions were based on extent of occurrence (EOO) mapping methods which erroneously indicate that a species is falsely present (error of commission) or falsely absent (errors of omission). Extent of occurrence maps are prone to commission errors as they intentionally ignore the internal extant of the area actually occupied by a species. In addition, it is suspected that historical inaccuracies may have been unwittingly adopted from researcher to researcher that perpetuated earlier omissions due to further insufficient ground-truthing.

2.8 CONSERVATION STATUS

Readily acknowledged as an uncommon to rare species (Shortridge, 1934:96;

Smithers, 1971:128; Visser, 1976:73; Stuart, 1982:8; Stuart & Wilson, 1988:23;

Nowell & Jackson, 1996:8; Sunquist & Sunquist, 2002:80; Skinner & Chimimba,

2005:406; Sliwa, 2013:204), the IUCN Red Data global classification lists BFC as

Vulnerable with a total effective population size of 10 000 mature breeding individuals and declining (Sliwa, 2008). It is ranked as Category 2 for global vulnerability and considered the most vulnerable of the sub-Saharan cat species by the Cat Specialist Group (Nowell & Jackson, 1996:7). The species has also been included as an endangered species since 1975 on Appendix 1 of the

Convention on International Trade in Endangered Species of Wild Fauna and

Flora (CITES). In the most recent update on the assessment carried out by Sliwa,

Wilson, Küsters and Tordiffe (in prep.) in 2014-15, the global IUCN conservation status has remained unchanged.

Nationally in South Africa, the species is listed in the Red Data Book only as Least

Concern (Friedman & Daly, 2004:172-173), but they are listed as a Protected

Species under the Biodiversity Act 10 of 2004 (SA, 2004) and still included in the

2007 updated species list. It is protected across most of its range (Nowell &

Jackson, 1996:9) excluding Namibia and Zimbabwe. The more recent conservation assessment, carried out by Wilson, Sliwa and Drouilly (in prep.) in

2014-15 for the updating of The Red List of Mammals of South Africa, Swaziland and Lesotho, has seen the status uplifted to Vulnerable based in part on the data presented in this dissertation.

Elsewhere in the southern African subregion, no other countries conduct their own conservation assessments, and adopt the listings established by the IUCN. As such, BFC are listed as Vulnerable in Botswana (Ministry of Environment, Wildlife and Tourism, 2009:16) but remains unlisted in Namibia (Ministry of Environment and Tourism, 2014) despite it being present in the region. Because the species is has been thought not to be present in the following regions, it is also gone unlisted in Angola (Ministry of Environment, 2009), Zimbabwe (Ministry of Environment &

Natural Resources Management, 2010), Zambia (Ministry of Tourism,

Environment and Natural Resources, 2009) and Mozambique (Ministry for the

Coordination of Environmental Affairs, 2014).

2.9 PRINCIPAL THREATS

Currently, there are a number of acknowledged threats to BFC all of which are human related (Nowell & Jackson, 1996:9). These include indiscriminate persecution and accidental deaths by farmers during poisoning and predator control activities such as spraying for locusts, and the hunting and trapping of other target species such as jackal and caracal. In addition, Nowell and Jackson

(1996:9) suggest that overgrazing by livestock that results in the loss of prey base through the degradation of the cats preferred habitats further threatens the BFC.

CHAPTER 3

RESEARCH METHODS

3.1 INTRODUCTION

Throughout the literature reviewed, the black-footed cat (BFC) (Felis nigripes), has been described as uncommon to rare (Shortridge, 1934:96; Smithers, 1971:128;

Visser, 1976:73; Stuart, 1982:8; Stuart & Wilson, 1988:23; Nowell & Jackson,

1996:8; Sunquist & Sunquist, 2002:80; Skinner & Chimimba, 2005:406; Sliwa,

2013:204). This was the basis for explaining the general lack of distributional data available for the species.

Using a variety of research techniques, historical and current geographical locality data was gathered for the time span of 1824 – 2014. Data collection began in

2006 and continued continuously for eight years, but is still ongoing on an ad hoc basis.

3.2 RESEARCH DESIGN

A combination of non-experimental research techniques were applied in this longitudinal study that relied on both qualitative and quantitative methods. The need for this type of integrated and interdisciplinary research has become widely recognised in situations where social science and natural science situations are intertwined (Nuijten, 2011:204). Qualitative aspects included the informal social interview processes during the citizen science surveys, whilst quantitative aspects

included the semi-structured formal interviews conducted by the researcher, published records and museum specimens.

3.3 DATA COLLECTION

Data collection of geographical locations for BFC began in 2004 as part of the overall investigation of the BFC by the Black-footed Cat Working Group (BFCWG), but became more formalised and structured in 2006. A number of key techniques were used to gather data for this study. Several methods overlapped more than one objective and/or key question/s, and some were used in conjunction with other methods.

3.3.1 Literature review

As indicated previously, historically the research effort has been extremely low for this species. The majority of historical records were handled using a retrospective systemic literature review and document analysis to identify, summarise, appraise and validate all geographical references. Sources of information were obtained from a wide variety of published sources including, but not limited to: travel journals (e.g. Burchell, 1824:592; Livingstone, 1857); published accounts of museum specimen holdings (e.g. Pocock, 1907:669-674; Shortridge, 1934:96); archaeological records (Klein, Crux-Uribe & Beaumont, 1991:99; Kuhn et al.,

2011:7); International Zoo Yearbooks; zoological necropsy reports; journal articles

(e.g. Lynch, 1975:109-139; Stuart, 1982:7-9; Watson, 2006:114), and reference guides (Smithers, 1971:392; Skinner & Smithers, 1990:419; Skinner & Chimimba,

2005:407). Searches were extended into grey literature to be more inclusive, and included unpublished and online reports (Joubert, Morsbach & Wallis, 1982:17;

Rowe-Rowe, 1992:23; Sliwa, Wilson, Lamberski & Herrick, 2007; Sliwa, Wilson,

Lamberski & Herrick, 2008; Sliwa, Wilson, Lamberski & Herrick, 2009a; Sliwa,

Wilson, Lamberski & Lawrenz, 2009b; Sliwa, Wilson, Lamberski & Lawrenz, 2010;

Sliwa, Wilson, Lamberski, Lawrenz & Herrick, 2011; Sliwa, Wilson, Lamberski &

Lawrenz, 2013; Sliwa, Wilson, Lamberski & Tordiffe, 2014; Sliwa, Wilson, Küsters,

Lawrenz, Eggers et al., 2015).

3.3.2 Museum records

Aside from published museum records available in referenced literature, all known national and international museums with African mammal collections were approached to obtain data of their BFC holdings. Data were sourced from six

United States of American museums namely: the American Museum of Natural

History3, California Academy of Sciences4, The Field Museum5, Natural History

Museum of Los Angeles County6, Museum of Vertebrate Zoology7, Smithsonian

National Museum of Natural History8. Data was also obtained from six South

3 www.amnh.org/our-research/vertebrate-zoology/mammalogy/database

4 www.calacademy.org/scientists/ornithology-mammalogy-collection

5 www.fieldmuseum.org/science/research/area/mammals

6 www.collections.nmnh.si.edu/search/mammals

7 www.mvz.berkeley.edu/Mammal_Collection.html

8 www.vertebrates.si.edu/mammals/mammals_databases.html

African collections namely the McGregor Museum9, Durban Natural Science

Museum10, Amathole Museum11, National Museum12, Iziko Museum13 and Ditsong

Museum14, and one Namibian museum the Natural History Museum15. Data was either sourced via email requests, visits to the collections or from web-based online archives.

3.3.3 Web-based resources

A number of free and open web-based resources were consulted for species occurrences. These focus on making scientific data sets on biodiversity available via the Internet. The data are provided by many institutions from around the world and these data are accessible and searchable through single entry portals.

International resources consulted were Global Biodiversity Information Facility

(GBIF)16, the International Union for Conservation of Nature (IUCN)17, World

Wildlife Fund (WWF)18 and Encyclopaedia of Life (EOL)19.

9 7 Atlas Street, Herlear, Kimberley,8301, South Africa

10 Anton Lembede Street, Durban, 4001, South Africa

11 Albert Road, King Williamstown, 5601, South Africa

12 36 Aliwal Street, Bloemfontein, 9301, South Africa

13 25 Queen Victoria Street, Cape Town, 8001, South Africa

14 432 Paul Kruger Street, Pretoria, 0001, South Africa

15 Robert Mugabe Avenue, Windhoek, Namibia

16 www.gbif.org/

17 www.iucn.org/

18 www.worldwildlife.org

Also consulted was the African citizen science project MammalMAP20 jointly run by the Animal Demography Unit (ADU) at the University of Cape Town and the

Mammal Research Institute (MRI) at the University of Pretoria. This is a digital database aimed at updating the distribution records of all African mammal species.

Incidental and random locality records were sourced by routinely carrying out open

Internet searches using “small-spotted cat”, “black-footed cat”, “Felis nigripes”,

“miershooptier”, “klein gekolde kat” in Google search engines. These positive search results were usually sightings or listed attractions on web pages of various guest farms and national park forums.

3.3.4 Observations

Observations records were divided up into two types. Those made by the researcher or other experienced researchers during field surveys, or those collected by solicitation from the public.

3.3.4.1 Field surveys

Faunal surveys during routine field work from 1988 to present, under the auspices of McGregor Museum zoological research and with the BFCWG contributed a number of direct observations of BFC using night time spotting techniques as described in Sliwa et al. (2007), Sliwa et al. (2008), Sliwa et al. (2009a), Sliwa et al. (2009b), Sliwa et al. (2010), Sliwa et al. (2011), Sliwa et al. (2013), Sliwa et al.

19 www.eol.org/

20 www.mammalmap.adu.org.za/

(2014) and Sliwa et al. (2015). A number of BFC were also captured and released after being fitted with telemetry collars for monitoring purposes. Other incidental observations included roadkill specimens, and skins from the collections of jackal hunters.

In rare instances, indirect observations of BFC included footprints and scats were accepted from experienced trackers. Various other indications such as habitat appropriateness, presences of sympatric species, and other appropriate bio- indicators were considered.

3.3.4.2 Citizen scientist records

Citizen science typically refers to research collaborations between researchers and members of the public or volunteers to expand opportunities for scientific data collection. This technique of involving the public to acquire information on a variety of wildlife species and wildlife-related issues is advantageous for the researcher. It maximises the amount of data collected with minimum time and effort, and hence the costs (Webster & DeStefano, 2004:70). Combinations of qualitative and quantitative techniques were used to allow for a more comprehensive understanding of the various data obtained during the interview process (Nuijten,

2011:198).

The citizen scientists varied from land owners and managers, conservation practitioners, farm employees, predator control officers, taxidermists, international shipping agents, private captive keepers (legal and illegal), professional hunters,

tourists and social media. Included here were field biologists and museum scientists with unrelated specialities.

The majority of records were solicited either by the use of specially designed posters in English (Annexure A) and Afrikaans (Annexure B) of which 500 each were distributed (including electronic versions) throughout South Africa, and to a limited extent to neighbouring countries of Namibia and Botswana.

Such solicited information has been used on similar projects: bobcats (Lynx rufus)

(Broman, Litvaitis, Ellingwood, Tate & Reed, 2014:232), Scottish wildcat (Felis silvestris silvestris) (Davis & Gray, 2010), Iberian lynx (Lynx pardinus) (Palma,

Beja & Rodrigues, 1999:814), African wild dogs (Lycaon pictus) (Lindsey, du Toit

& Mills, 2004:144) and greater roadrunners (Geococcyx califnornianus) (Webster

& DeStefano, 2004:70).

Contributions were also received in response to raised awareness of the species after seeing the logoed BFCWG project field vehicle, magazine and newspaper articles, specific website requests on the Internet portals of the International

Society for Endangered Cats (ISEC)21, Wild Cats22, Cat Conservation Trust23, and

21 www.wildcatconservation.org/black-footed-cat-project/

22 www.black-footed-cat.wild-cat.org

23 www.karoocats.org/Distribution.aspx

social media pages such as Save the Black-footed Cat!24. Specific Internet predator control and hunting forums25 were also targeted for information. The nature of citizen scientist records included historical and current records of sightings, roadkill specimens, hunted (accidental and deliberate), trapped animals and camera trap photos.

Records were then followed up using semi-structured semi-formal interviews

(telephonic, face-to-face, correspondence and online chat groups). These interviews consisted of closed questions (name, contact details, locality details, date) and open questions (observational details about the sighting including description of the animal seen, quality of the sighting, weather conditions, habitat, predator control activities in the area, behaviour of the cat, number of current and past sightings and sympatric species questions). Datum was recorded on a basic form (Annexure C).

Hearsay records were also collated from data collected on behalf of the researcher by third parties. Such data was routinely collected by Endangered Wildlife Trust26

(EWT) officials and conservation practitioners at annual agricultural shows (e.g. annual NAMPO Harvest Day27) and at district agricultural meetings and during the

24 www.facebook.com/groups/9242632404/?fref=ts; www.facebook.com/pages/Black-footed-Cats/261631538251?fref=ts, and www.facebook.com/groups/39518046010236?fref=ts.

25 Example: www.jaracal.com/index.php

26 www.ewt.org.za

27 www.nampo.co.za 31

course of field surveys and routine investigations. This data was mostly limited to absence and/or presence on a specific property with no additional details. Where contact details were supplied, follow-up interviews were conducted by the researcher.

Other incidental sources of datum were also obtained from the Northern Cape

Department of Environment and Nature Conservation and the South African Police

Services investigative and prosecution case files.

3.4 DATA ANALYSIS

Data analysis involved various steps, from the capturing and grading of records followed by the mapping of data localities. Thereafter, temporal and spatial analysis was applied to the record data whilst GIS analysis was carried out on the mapping data.

3.4.1 Geographical records

All data records received were recorded in a specifically-designed Microsoft

Access database software which made provision for transfer to a Microsoft Excel spreadsheet for statistical analysis. Each record received a unique reference code

(observation number) to cross-reference records and to maintain anonymity where requested. Where possible, each record contained the following details: 1) record details; 2) record type; 3) sex; 4) age; 5) date; 6) locality details including nearest town, province, country; 7) quarter degree square (QDS) and coordinates in

degrees and minutes; 8) whether or not the locality was calculated; 9) observer rating; 10) name and contact details of the observer/collection/data source; 11) additional details relating to the record; and 12) habitat details.

Without clear guidelines, temporal data was divided into two categories. Historical data were considered as all records prior to 1990, and current data were considered all records from 1990 onwards. The significance of using 1990 as the inflection was based on research on the species beginning in earnest by Sliwa

(2004, 2006) and assisted by the researcher from 1992-1998. This sparked increased research effort and interest in the species. In addition, precision locality instruments such as global positioning systems (GPS) became more readily available allowing for more accurate geographical recording.

Due to the limited number of historical records, there was some duplication of data

(sightings and museum records) in instances when museum institutions had made their data available to more than one online resource, or when a journal publication referenced a specific specimen in a museum collection. These duplicated records were identified and removed.

Locality data was recorded to point data but only to degrees and minutes level and indicated as “locality uncalculated” when exact locality was provided from GPS coordinates. In instances of only farm name, or distance and direction from nearest town being available, the location was calculated using Garmin

MapSource28 and Garmin BaseCamp29 topographic maps and online world gazetteers30.

Records received were assessed using identifying physical characteristics, geographical location and other supplied details. This was particularly important with records from citizen scientists who varied greatly in their knowledge of the species and attention to detail. Since it was not possible to measure implicitly the reliability and validity of the data, only credible records were included in the database, and all were evaluated and categorised with an observer rating (OR) based on the quality of the record:

 Observer Rating 1 (OR1): single, unconfirmed record (no photograph or

supporting evidence); identity may be doubtful; or although record was

considered reliable, precision locality data unavailable; observer credibility

unknown.

 Observer Rating 2 (OR2): single, confirmed record (with supporting evidence,

tangible, photographic or descriptive) and locality data available; observer

credible.

 Observer Rating 3 (OR3): multiple, confirmed records for a specific QDS.

28 www.garmin-mapsource.en.lo4d.com

29 www.garmin.com/en-ZA/shop/downloads/basecamp

30 Example: www.gazetteering.com/africa/

3.4.2 Mapping

No digital historical geographical distribution maps were available and these had to be digitised from scanned published reports in order to generate free-hand traced polygons. The graphic files were then uploaded to online software Map

Warper31 and georecitified. All data and georecitified images were imported into

Quantum GIS 1.8.0-Lisboa (QGIS, 2015)32 for GIS analysis. All resulting maps were generated using ESRI (ESRI, 2011) and QGIS shapefiles and a coordinate reference system (CRS) with a WGS 84 projection. Statistical analysis within

QGIS was used to determine range sizes and answer other mapping queries.

Geographical distribution (extent of occurrence) maps were created using the marginal occurrences method (Gaston & Fuller, 2009:2) which uses the outermost locality records of a species and interpolating between these to delineate the boundary. Obvious geographical features known to be unsuitable habitat for the species (i.e. desert with <200 mm of rainfall, mountains, areas above 2 000 m above sea level, large bodies of water, etc.) were avoided.

Area of occupancy (AOO) was calculated using QDS grid cells. This spatial scale size was chosen because data was collected and averaged to QDS level only, and because it the method used in the MammalMAP project. A cell in this grid in the southern hemisphere is about 685 km2 and this is considerably less than the 50 km buffering radius applied to each record that is approximately 8 000 km2 in extent.

31 www.mapwarper.net

32 http://qgis.osgeo.org

3.4.3 Statistical method

All statistical analysis (Pearson correlations and trends) were calculated in

Microsoft Excel using Analysis Toolpak. Various built-in vector analysis, geoprocessing and spatial query tools in QGIS Version 1.8.0 ‘Lisboa’ (2015) were used during the initial mapping process to answer basic mapping queries.

3.4.4 Population size estimates

In this investigation, an isopleth (heatmap) was created to estimate population size based on the methods recently applied to BFC conservation assessment in The

Red List of Mammals of South Africa, Swaziland and Lesotho (Wilson, Sliwa &

Drouilly, in prep.) currently under revision. However, this mapping exercise was extended to include the entire southern African subregion rather than just South

Africa, Swaziland and Lesotho and included both historical and current data.

Basic point map data created in the mapping exercise was imported into ArcGIS®

(ESRI, 2011)33. The Kernel Density tool in ArcGIS was used to estimate kernel densities around BFC records. The densities were converted to isopleths containing 30, 40, 50, 60, 70, 80, 90 and 95% of the sightings. Isopleth lines were converted to polygons and the area of each polygon was calculated, after the polygons were projected to Africa Albers Equal Area Conic projection.

33 http://www.esri.com/software/arcgis/new 36

High density clusters of BFC localities were assumed to contain 50% of sightings

(i.e. the 0.5 isopleth), whereas medium-density clusters were assumed to contain the next 20% of sightings (the 0.7 isopleth) while low density areas covered the remaining 25% of sightings (0.95 isopleth).

The area (in km2) was then calculated for each of the densities. For South Africa, the kernel density polygons were clipped to the national boundary. Densities of cats in the different areas were estimated at 0.03, 0.02 and 0.01 per km2 (Sliwa,

2004:100; Sliwa et al., 2007; Sliwa et al., 2008; Sliwa et al., 2009a; Sliwa et al.,

2009b; Sliwa et al., 2010; Sliwa et al., 2011; Sliwa et al., 2013; Sliwa et al., 2014;

Sliwa et al., 2015). The three density zones were summed, and this total was adjusted to 70% to represent the mature population structure. The total areas for high, medium and low density clusters were calculated and the corresponding population totals estimated.

3.5 LIMITATIONS

All studies have limitations, some of which are related to methodology or survey design whilst others are related to the number or types of data collected or the procedures used. Those with the greatest impact on the findings and the ability of these to answer the research questions are identified here. The use of the semi- structured follow-up interviews and rigid application of an observer rating to each record was designed to address and minimise the impacts of the following limitations:

3.5.1 Observer biases

According to Durheim (2006:129) there are two important factors to take into account when evaluating a quantitative survey. The first is reliability, the extent to which a survey produces similar results on repeated surveys i.e. this is a measure of consistency involving equivalence, stability and internal consistency.

Equivalence refers to the degree to which the various survey methods collected the same data. Stability refers to whether or not another researcher would achieve the same results with similar, repeated surveys. Internal consistency refers to the degree to which the various aspects of the data collected for each record relates to the record as a whole.

The second factor is validity, the extent to which the instrument measures what it is supposed to measure i.e. whether or not it supports the data and whether this is relevant to the research questions. Data can be reliable but this does not necessary make it valid! Of relevance here, is the predictive validity of the data i.e. whether is logically accurate to predict the presence of the species based on the record, and the content validity that measures the extent to which each data record contained all of the necessary details in order to achieve an accurate assessment.

In many cases, citizen scientist records were incidental and opportunistic observations. The data were collected without standardised field protocols and without explicit sampling design. Such anecdotal and opportunistic occurrence data and inconclusive physical data are often used to assess the current and

historical ranges of rare or elusive species and can be fraught with problems unless strict standards are applied (McKelvey, Aubry & Swartz, 2008:549). It is a challenge to achieve reliable estimates of distribution trends because this data differs in terms of field efforts over time (observation bias), from incomplete and selective recording (reporting bias), and from geographical bias.

Detection bias was also a factor given that the species is considered naturally rare, is very small, cryptic, nocturnal. It has several colloquial names and often mistakenly thought to be, or misidentified as another species e.g. genets (Genetta spp.) and juvenile African wildcats.

Other influences can also include education and interest levels, as well as confusion over the name of species. Truthfulness due to fear of persecution by conservation authorities when records were linked to a contravention of provincial or national legislation (e.g. being in possession of a wild-caught animal without a permit, or having hunted an animal either deliberately or accidentally) may also have been a factor. This is known as the Hawthorne Effect (McBride, 2013:89) and occurs when respondents change their behaviour or claim a different behaviour because they are being questioned and perceived the interviewer to be associated with, in this case, conservation agencies. The effect was minimised by reassurance during the personalised interview process that information was to be treated strictly confidentially.

This aim of collecting data during the interview processes by using a specific set of questions was designed to improve reliability. The questions tested the

interviewee’s knowledge and attitudes to the species and other factors that might be related to the presence or absence of the BFC. This method also allowed for additional data regarding habitat, weather conditions, predator control activities in the area, behaviour of the cats, and was used to assess the validity of the observation based on the trustworthiness of the responses in terms of credibility and/or dependability.

3.5.2 Historical imprecision

Verification of the historical records was not always possible, even from museum specimens that are still present in collections. All specimens in southern African collections were examined, but for logistical and practical reasons it was not possible to examine material held in international collections.

The majority of historical locality names that have been changed or are now spelt differently and therefore these necessitated additional georeferencing.

Furthermore, localities or provenance tended to be vague or imprecise either due to poor reporting in the literature or some uncertainty by the reporter. These then had to be calculated to an averaged degree and minute position because precision coordinate recording devices such as GPS devices were not available prior to

1990. Boshoff and Kerley (2010:32) also found that 82% of written historical records were useful for compiling historical distribution records despite some of them dating back to 1750, and thus historical records are still considered useful, particularly if they have been evaluated for reliability.

3.5.3 Record verification

As previously mentioned, citizen scientist records were often incidental and opportunistic observations. Information can be erroneous – misidentification as genets and juvenile African wild cat being the most common problem. These were however eliminated through questioning and reviewing of photographic evidence.

With referenced literature, museum records and web-based biodiversity data it was not possible to confirm the correct identification of the animal cited unless, for example, the record or reference was accompanied by a photograph.

Obvious inaccuracies in record details (e.g. exceptional tail lengths or body weights and sizes, etc.) which indicated a high probability of the specimen being another cat species or a genet were noted in several cases when reviewing morphological data. Such records were discarded from the database and from the subsequent analysis.

Species identifications in virtual museums such as MammalMAP were carried out by a third party panel of mammal experts appointed by the Animal Demography

Unit34, and were deemed reliable records.

3.5.4 Gaps in the data

Given that the geographical distribution of the species was expected to be throughout southern Africa, it was obviously impossible to physically survey the

34 http://adu.org.za/index.php 41

entire region in which the geographical range of the species is thought to occur during this the research period. A targeted survey was conducted, but as with all survey samples, a total survey error (the difference between the total number of records and the estimate of the parameter based on the sample survey) is to be considered. Many areas falling within the known or suspected geographical range were not investigated due to time and logistical constraints, and with landowners being unaware that data on the species was being collected.

Many international and national museums have significant backlogs in data capturing (Vollmar, Macklin & Ford, 2010:104). The resulting unaccessioned

(uncatalogued) specimen data was thus unavailable for inclusion in this dissertation.

3.6 ETHICAL CONSIDERATIONS

As is common practice with sensitive datasets, all data were modified and coded to ensure the confidentiality and anonymity of the individual records and the source of the data. Each record was given a unique record number for referencing purposes. The precise localities will not be made publicly available.

However, given the potential importance of the species from a future historical viewpoint and for conservation reasons, a full hardcopy of the database will be placed in the document archives at the McGregor Museum, Kimberley, South

Africa, with a 20-year embargo. This will ensure that the data sources will not be

published or made available until a certain date or certain conditions have been met.

Ethical clearance for the project was granted by the Research Ethics Committee of the Tshwane University of Technology in October 2010 (Annexure D).

3.7 DISSERTATION OVERVIEW

The combined results and discussions will follow in two separate chapters, each covering one of the main key questions. Chapter 4 will focus on the geographical distribution of BFC, whilst Chapter 5 will focus on the conservation status of BFC.

The final chapter with provide a synthesis of the findings and conservation implications that will then lead into suggested management recommendations for the species.

CHAPTER 4

RESULTS AND DISCUSSION: GEOGRAPHICAL DISTRIBUTION

4.1 INTRODUCTION

A population’s geographical distribution is essentially a measurement of the range of a species in a region, area, or specific locality. Knowing a species’ geographic range and habitat preferences is essential to conservation efforts and the developing of management plans.

The historical paucity of data in general on the black-footed cat (BFC) (Felis nigripes) has led to inconsistencies and perpetuated inaccuracies in current literature that in turn has affected the accuracy of conservation measures. The naturally rare, cryptic appearance, small size and secretive nature of BFC has contributed considerably to this data deficient situation.

This research investigates the accuracy of previous geographical distribution estimates for the species and attempts to propose an updated version of their geographical distribution map. This new current geographical range was compared to the historical ranges to determine the reasons for the different estimates over the past 40 years, and updates the geographical distribution map of the BFC.

4.2 REVIEW OF SURVEY RESULTS

Survey results were examined to determine the types of records and the frequency and accuracy of these records. The spatial and temporal distribution of records was also examined.

4.2.1 Record and observer categories

More than 887 records were investigated. Obvious erroneous records of similar species such as genets (Genetta spp.), African wildcat, domestic cats (Felis silvestris catus) and young serval (Leptailurus serval), cheetah (Acinonyx jubatus) and leopard (Panthera pardus) were eliminated and all duplicate records were removed. A final 790 records were used for data analysis. All records were verified and given an observer rating (OR) category. These records were clustered into various record types (Figure 4.1).

The majority of the total survey records (73%, n = 577) were categorised as OR 2

(single, confirmed observations). The second highest category (14.6%, n = 115) was OR 3 records (multiple, confirmed observations in a particular area QDS), followed by OR 1 records (12.4%, n = 98) which were doubtful or unconfirmed records. These results suggested that 87.6% of the data were reliable and valid.

600

500

400

300 n = 790

Number of records of Number 200

100

0 Sight Capture- Specimens Captive Roadkill Fossil Records Release OR 3 71 31 3 8 2 0 OR 2 421 117 15 9 13 2 OR 1 61 33 0 1 3 0 Type of record and observer rating

FIGURE 4.1: Black-footed cat survey results grouped according to type of record and observer rating categories where OR 1 is unreliable record; OR 2 is a single, reliable record, and OR 3 is are multiple confirmed records for a specific quarter degree square

4.2.1.1 Sight records

Sight records (n = 553) accounted for the greatest number of records (70%) in the survey. These records were mostly contributed directly to the researcher by members of the public, farmers, professional hunters, conservation field officers and other biologists. Other records were indirectly collated by the researcher using

published sightings, Internet searches, citizen science projects (e.g. MammalMAP) and specialised predator and hunting social platforms such as www.jackalpro.com.

In addition, sightings were recorded during the course of BFCWG field studies and personal investigations.

The nature of these sight records included physical observations of black-footed cats; evidence of cats such as spoor or scats; photographic or video evidence; journal accounts; grey literature, and accounts of deliberately or accidentally killed animals (victims of humans or dogs) that were not formally collected as a specimen for legal reasons. Sightings of BFC were made in various situations including, but not limited to, a variety of natural habitats, sheep farming areas, crossing the road, in agricultural fields and around homesteads. Of the sight records, 76.1% were category OR 2, whilst 12.8% were OR 3 and 11% were (OR

1). This meant the majority (88.9%, n = 492) were reliable records.

4.2.1.2 Specimen records

Specimens records (n = 181) accounted for the second highest proportion of survey records at 22.9%. These records included taxidermist client records

(hunting registers) of mounted animals, and museum records of accessioned skin, skull and/or skeleton specimens as well as skin samples taken from privately owned skins.

The majority of these records (81.8%, n = 148) had good provenance (locality) data as could be expected from museum specimens and hunting registers, whilst

the remaining records had only generalised information (e.g. a country or province name only) that did not allow for localities to be calculated.

4.2.1.3 Captive records

Captive records (n = 18) accounted for a small proportion (2.3%) of the total records and included animals that were currently in captivity (legally or illegally) in

South Africa or elsewhere that had known provenance (OR 2 and/or 3). Other individual captive BFC were known to exist but either were captive-bred or the original capture location unknown and these were not included in the survey.

4.2.1.4 Capture-release records

Capture-release records (n = 18) accounted for a small proportion (2.3%) of the total records and included selected animals that were handled by the BFCWG at various locations during field surveys, as well as a number of other records in which cats were unintentionally caught in non-lethal traps (Figure 4.2) and released elsewhere.

To prevent over-representation in the survey, only specific localities in which the

BFCWG has conducted fieldwork were included. All the records had good provenance data with the exception of one record in which the capture details were not disclosed by the parties involved.

FIGURE 4.2: A black-footed cat unintentionally trapped by a farmer in a cage trap set for larger problem-causing predators which was later released after being correctly identified

4.2.1.5 Roadkill records

Only 2.3% (n = 18) roadkill records were correctly reported to or directly observed

(and collected as museum specimens) by the researcher. Black-footed cats were roadkill victims on tar roads as well as on gravel roads. Of these, three records had inexact locality details, or the accompanying photographic evidence could not be definitively identified as the correct cat species. A number of the observations submitted were misidentifications, usually African wildcats or genets. However, the

BFCWG has documented cats expertly negotiating busy roadways (Figure 4.3).

FIGURE 4.3: A sequence of photographs showing a male black-footed cat (BFCWG research subject “Kubu”) deliberating and successfully negotiating busy traffic on a national road (N8) near Kimberley, Northern Cape, South Africa (Photograph: A. Sliwa)

4.2.1.6 Fossil records

Prior to this study, it was thought that no fossil records for this species existed.

Literature searches revealed two fossil records from two different localities namely

Equus Cave near Taung, North West Province and Malapa Hominin site near

Krugersdorp, Gauteng (Figure 4.4) in South Africa. These records were included in the survey as they assist in establishing an evolutionary timeline for the species.

FIGURE 4.4: A lateral view of a black-footed cat jaw fragment (UW 88-517) recovered from Malapa Hominin site, South Africa (Kuhn et al., 2011:7)

4.2.2 Spatial distribution of records

Records from six southern African countries were included in the survey (Figure

4.5). Of these records, 344 (43.5%) had exact coordinate locations recorded, whilst 409 (51.8%) had sufficient locality information that a proximate (degrees and minutes) location could be determined. Only 37 (4.6%) had no location data or data that only placed the record to a province or country.

Of the six southern African countries, only four (0.5%) incidental records came from countries (Angola, Zimbabwe and Zambia) beyond the current known geographical range of which the later record from Zambia is highly doubtful and the historical Angolan records warranting further investigation. The Zimbabwean record was considered valid.

South Africa Zimbabwe 0% Zambia 0% 90% Unknown 3% Angola 0%

Botswana 4%

Namibia 3%

FIGURE 4.5: Spatial distribution of black-footed cat survey results grouped by country

A significant majority of records (90%, n = 706) were South African, whilst

Botswana and Namibian records accounting for 4.3% and 3.3% respectively.

These were largely historical records. However, there was considerable underreporting of historical records in Namibia. In a survey conducted in 1982

(Joubert, Morsbach & Wallis, 1982) of 2 433 farms in 12 different districts, 306

(12.6%) reported the presence of BFC on the property. Reporting rates varied per district from 1.5% to 30.4% (µ = 12.6%). However, with the data having been grouped to a district level, it was not possible to plot individual occurrences. Each district was assigned a OR 2 only despite multiple records. This was because no exact localities could be determined and there were no clear indications how reliable the records were, or how they were evaluated.

No records were obtained for Swaziland, Lesotho or Mozambique, and with South

African data forming the majority of the records, it was worth investigating the spread of records within the nine official provinces of the country which are represented in Figure 4.6.

Eastern Cape Western Cape 15% Gauteng 32% 1% KwaZulu-Natal 2%

Free State 14%

Limpopo 2% North West 4% Mpumalanga 4% Northern Cape Unknown 25% 1%

FIGURE 4.6: Spatial distribution of black-footed cat survey results grouped by South African provinces

Of the provincial records, 86% were from localities in the Northern Cape, the

Eastern Cape, Western Cape and the Free State. The remaining provinces North

West, Limpopo, Gauteng and KwaZulu-Natal shared the other 13% of the records with only six records having unknown South African provenance.

4.2.3 Temporal distribution of records

The 790 survey records were divided into historical records that consisted of all records up to until the end of 1989. Current records were considered all records from 1990 onwards. Seventeen records had periods that spanned more than one decade resulting in 813 temporal records. Periods were then grouped into decades dating back to 1900 with no gaps. Due to the scarcity of records (n = 6)

from the 1800s, these were grouped together, as were the two fossil records

(Figure 4.7).

350 Fossil & historical records 311 300 Current records

Record date unknown 250 234

200

150

100 81 57

50 35 39 Number of records pergrouped years records of Number 10 17 2 6 7 1 3 2 8 0

Grouped Years

FIGURE 4.7: Temporal distribution of black-footed cat distribution records

There were 194 historical records spanning 11 periods. For the remaining three current periods, 602 records were documented, with the majority falling in the last decade and a half following the initiation of this study.

The remaining 17 records had no data and although were all historical records in overseas museums accompanying specimen data could not place the records in

either the 1800s or 1900s. Since either provenance or morphological data was available, these records remained in the database.

Altogether, there were more temporal records than actual records due to 23 records of the original 790 reporting data spanning more than one decade. This was more apparent in the current records and confounded the mapping statistics later.

4.3 REVIEW OF HISTORICAL MAPS

An extensive literature review revealed only seven authoritative papers or references in the past 35 years specifically addressing the envisaged geographical distribution of BFC, many of which contained only unrefined range maps or referenced each other (Stuart, 1982:8; Stuart & Stuart, 1988:152; Smithers,

1983:392; Skinner & Smithers, 1990:419; Friedman & Daly, 2004:173; Skinner &

Chimimba, 2005:407; Wilson & Sliwa, 2007). Digitised versions can be seen in

Figures 4.8 to 4.12 and the approximate area (km2) of geographical distribution calculated for each. In each instance, the assumption is that the EOO was established by mapping the marginal occurrences and ignoring outlier records.

FIGURE 4.8: Black-footed cat distribution (approximately 1 236 178 km2) as presented in Stuart (1982:8)

FIGURE 4.9: Black-footed cat geographical distribution (approximately 1 253 534 km2) as presented in Smithers (1983:392)

FIGURE 4.10: Black-footed cat geographical distribution (approximately 1 401 004 km2) as presented in Skinner and Smithers (1990:419)

FIGURE 4.11: Black-footed cat geographical distribution (approximately 1 530 437 km2) as presented in Skinner and Chimimba (2005:407)

FIGURE 4.12: Black-footed cat geographical distribution (approximately 2 043 713 km2) according to Wilson and Sliwa (2007)

Between 1982 and 2008, there has been a consistent (y = 189197x + 925381, R2

= 0.82) and significant (Pearson correlation: r = 0.88, n = 5) expansion in the probable geographic distribution represented in publications as seen in Figure

4.13. The range extensions appeared largely in a northerly and easterly direction whilst the southern and western range limits remained mostly unchanged.

2500

2000 y = 189197x + 925381 R² = 0.8202 1500

1000

500 Area km2 1 = 000 Area 0 Stuart 1982 Smithers Skinner & Skinner & Wilson & 1983 Smithers Chimimba Sliwa 2007 1990 2005 Publications

FIGURE 4.13: Changes in the area size (km2) of the envisaged black-footed cat geographical distribution as described in publications from 1982 to 2007

Stuart (1982:8) and Smithers (1983:392) both considered the majority of the range of BFC to be within South Africa (excluding Mpumalanga, Gauteng, Limpopo and

KwaZulu-Natal), with extensions into the eastern areas of Namibia and western, southern and central regions of Botswana. Although virtually identical in extent,

Smithers’ map suggests a slight range increase of 1.4%. Skinner and Smithers

(1990:419) again predicated a similar range, but increased the range north- eastwards Namibia to stretch marginally into far western Zimbabwe. This was the first account of the possible inclusion of Zimbabwe in the range of BFC. This range extension represented a 10.5% increase on Smithers’ 1983 estimates.

Fifteen years later, Skinner and Chimimba (2005:407) during a major revision of

Smithers’ work, accepted the previous range but made a narrow range extension along the borders between Botswana and Zimbabwe following through to between

Zimbabwe and South Africa. A small range extension to the range on the north- western section in Namibia was made too, resulting in an overall 6.8% increase to the previous authoritative estimate.

In 2007, Wilson and Sliwa suggested an extensive range increase to the eastern part of the range in South Africa and Botswana, but also northwards in Botswana and Namibia extending marginally into the extreme southern parts of Angola. This range extension was 26.4% range extension from that of Skinner and Chimimba

(2005:407), and a 39.5% increase on Stuart’s 1982 range estimation.

The total area of concurrence in all five maps was a central region of about

1 136 526 km2. This area made up 91.1% of Stuart’s 1982 original map but only

55.6% of the Wilson and Sliwa’s 2007 predicted range map.

4.4 CURRENT MAPPING

Mapping was separated by observer rating types, researcher and time period and compared. A new, improved geographical map was created and compared to previous mapping efforts.

4.4.1 Observer rating (OR) maps

From the 790 records, 754 records had distribution points that could be plotted. Of these records, 344 (45.6%) had precise coordinate localities, whilst 409 (54.2%) had localities that had to be calculated to the relevant averaged degrees and minutes based on the locality description.

Of these data points, 69 (9.2%) had a low reliability (OR 1) (Figure 4.14). The records were evenly scattered over the entire subregion and included points in

Angola, Zambia, Botswana, Namibia and South Africa.

FIGURE 4.14: Black-footed cat records with an unreliable record rating (OR 1) indicated as red points

Many of these records were considered valid sightings, but because a precise

QDS could not be calculated, they were rated OR 1. An example of this would be 61

“10km from Cradock”. The locality would then be calculated to the QDS in which

Cradock falls but in reality, the location may have fallen just outside this QDS.

Some 570 (75.6%) records were reliable and confirmed records, and categorised as OR 2 (Figure 4.15). The spread of the records included significant areas in

Namibia and Botswana, with a single record on the Botswana-Zimbabwe border.

Records in South Africa were concentrated in a band extending from the central regions down into the north-western parts of the Eastern Cape and the north- eastern sections of the Western Cape. There was a sparse scattering of records outside of the central regions gradually thinning out towards the country’s boundaries. There were no records from eastern KwaZulu-Natal southwards to the

Eastern Cape.

FIGURE 4.15: Black-footed cat records with a single, reliable rating (OR 2) indicated as blue points

The remaining 115 (15.3%) records were multiple confirmed records for a particular QDS (Figure 4.16). Whilst there were a small number of records in

Botswana, the majority were recorded in central South Africa, gradually thinning southwards into the Eastern and Western Cape, and elsewhere in the Free State and more northern regions of the country.

FIGURE 4.16: Black-footed cat records with a multiple, reliable rating (OR 3) indicated as black points

By overlaying all three observer rating maps (Figure 4.17) the OR 3 records fall within those of OR 2 records. However, there are a number of OR 1 records that fall outside the combined OR 2 and OR 3 records, particularly in the northern reaches of the range (Angola, Zambia and Namibia); to the east in South Africa in

Limpopo, and Mpumalanga reaching southwards into KwaZulu-Natal.

There were no records reported for within the boundaries of Lesotho, Swaziland or

Mozambique.

FIGURE 4.17: Combined black-footed cat observer rating distribution records (with unreliable OR 1 indicated as red points; single but reliable OR 2 indicated as blue points, and multiple and reliable OR 3 indicated as black points)

4.4.2 Historical records (up until the end of 1989)

There were 613 time period records, of which 193 were historical records that could be mapped (Figure 4.18). Records were distributed across Namibia,

Botswana and South Africa. Although evenly spread across the range, there appears to be a slight clusters of records from central South Africa and in the south-eastern region of the Eastern Cape.

FIGURE 4.18: Historical (up until the end of 1989) black-footed cat distribution records indicated as purple triangles

When the historical distribution maps (1982-1990) are overlaid for these records, it becomes apparent that the maps were not inclusive of all historical data. Figure

4.19 over-lays the three earliest maps (Stuart, 1982; Smithers, 1983; Skinner &

Smithers, 1990). Stuart’s 1982 map covered 155 (80.3%) and Smithers’ 1983 map

143 (74%) of the historical records represented in this map. This was expected to be slightly low, as their information would not have included records up until 1990.

Skinner and Smithers’ 1990 map covered 153 (79.2%) records. The majority of records excluded in all the coverages were on the edges of the distribution maps.

Records north- and westwards in Namibia were excluded, as were northern and south-eastern records in Botswana. South African records not included were from

Limpopo, Gauteng and Mpumalanga provinces, whilst a scattering of records from the Eastern Cape appear overlooked.

FIGURE 4.19: Composite map of black-footed cat historical (up until the end of 1989) geographical distribution with Stuart (1982) in pink; Smithers (1983) in green, and Skinner and Smithers (1990) in purple

Figure 4.20 over-lays the two later historical maps. The 2005 map of Skinner and

Chimimba covered 159 (82.4%) of the historical records whilst Wilson and Sliwa’s

2007 map covered 185 (95.9%) of the records. Skinner and Chimimba’s map followed a similar data omission as the other three historical distribution maps.

Wilson and Sliwa’s 2007 map was the most inclusive but overlooked some eastern records in South Africa.

FIGURE 4.20: A composite map of black-footed cat historical (up until the end of 1989) geographical distribution with Skinner and Chimimba (2005) in deep yellow, and Wilson and Sliwa (2007) in pale yellow

None of the researchers included either Swaziland or Lesotho in the distribution maps.

4.4.3 Current records (1990 onwards)

Of the 623 time period records, there were 589 current records that could be mapped (Figure 4.21). Only eight of these records were at the same location/s as historical records. The records span 14 years of which 2006-2014 saw concentrated efforts to document the distribution of BFC.

FIGURE 4.21: Current (from 1990 onwards) black-footed cat distribution records indicated as green diamonds

This data produced the first records for the far northern areas of Namibia and for the Okavango Delta. Two Angolan records from the early 1990s were discovered in literature, also the first records for this country. Two other observation records were reported, one from Zambia and the other from Zimbabwe near the Botswana border in Hwange National Park. Additional distribution range extensions were recorded into Limpopo, Mpumalanga, KwaZulu-Natal and deeper southwards into the Western Cape.

Virtually no recent records were reported to the researcher, recorded in literature, or acquired for museums in central Botswana or the central and western regions of

Namibia during this period. There were no records reported for Lesotho,

Swaziland or Mozambique.

4.4.4 Current geographical distribution of BFC

Using the combined observer rating map in Figure 4.17, a proposed EOO or geographical distribution was predicted in a manner similar to previous map techniques based on expert opinion and by mapping marginal occurrence records and ignoring vagrant outlier and dubious records (Figure 4.22).

The borderline (dashed green line) was created in Google Earth in order to avoid obvious geographical features known to be unsuitable habitat. Known unreliable records (e.g. Zambian, KwaZulu-Natal, and southern African coastline data points) and historically outdated records (e.g. near Cape Town) were also excluded.

FIGURE 4.22: Combined black-footed cat observer rating distribution records (with unreliable OR 1 indicated as red points; single but reliable OR 2 indicated as blue points, and multiple and reliable OR 3 indicated as black points) with a proposed current geographical distribution outline overlaid as a dashed green line

The resulting BFC geographical map (Figure 4.23) is similar in appearance to the historical maps. The area covered by the polygon is approximately 2 214 276 km2 in extent. This is 9.2% larger than the 2007 map of Wilson and Sliwa. The probable range expansion for the BFC continues to increase marginally as with previously reported distribution ranges and the trend of changes remains consistent (y= 21156x + 874144, R2 = 8968) and significant (Pearson correlation: r

= 0.9, n = 6). This range estimate is nearly twice of that proposed by Stuart in

1982.

FIGURE 4.23: Current proposed basic black-footed cat geographical distribution map (approximately 2 214 267 km2)

4.5 DISCUSSION

Arising from the data review, several limitations and assumptions should be highlighted and explained in greater depth. Further to this, the impact of the applied technique to produce a more accurate distribution and the implied findings are discussed.

4.5.1 Limitations of data and some assumptions

In studies such as these, there are often confounding factors that make it difficult to make comparisons and/or to infer temporal or spatial changes. However, the geographical distribution records and accompanying information presented here are likely to constitute the largest collated database of information on BFC to date.

Some confounding factors that had to be considered included:

4.5.1.1 Observer effects

The most obvious limitation in this research design was the incorporation of citizen science contributions. The benefits and risks of using crowd-sourced citizen scientist data has been critiqued by many authors since it become a mainstream ecological research tool (Dickson, Zuckerberg & Bonter, 2010:162), but the benefits of enlisting the public to collect large quantities of data across an array of habitats and situations over time are invaluable to scientists (Booney, Cooper,

Dickinson, Kelling, Phillips et al., 2009:977).

Many of the problems related to uncertainty and errors contained in sighting data, which formed a large proportion of the database. This is a common problem experienced by researchers on similar projects (Palma, Beja & Rodrigues,

1999:819; Lindsey, du Toit & Mills, 2004:144; Webster & DeStefano, 2004:71;

Broman et al., 2014:232), but during this research were eliminated by carefully checking for consistency and accuracy in the reports. The criteria used to determine the reliability of the records were probably overly conservative and resulted in about 90 records being rejected.

4.5.1.2 Data gaps

Precision of locality records without accompanying GPS coordinates could not be guaranteed and these records formed a large proportion of the database.

Localities were derived from averaged applicable degrees and minutes based on the locality description provided. However, this mapping imprecision was not considered significant as no locality could have been plotted further than one QDS away in error. Since a QDS is only 25 x 27.4 km in size, this is significantly less than the dispersal capabilities of a BFC estimated to be in a radius of 50 km

(Wilson, Sliwa & Drouilly, in prep.).

Verification of international museum specimens was also not possible, and several museums approached in Europe with known specimens did not respond. Some of the older accounts may also have been missed as they were unavailable online or difficult to locate as hard copies.

Known gaps that were of particular concern were those from Namibia due to the clustering of 306 records into only 12 (OR 2) records due to a lack of specific locality details. Essentially, this resulted in the loss of 294 individual locations.

It is also suspected that, in areas where knowledge about the species or where the research objectives of this survey were unknown, the presence of the species has gone unreported by citizen scientists. This is particularly evident in the countries outside of South Africa.

Even in regions where there was knowledge of the project, reporting may have been hampered by other factors such as the elusive nature of the cat and its nocturnal nature. Thus, animals that occurred in less accessible areas or farmers/hunters that did not move about at night would have overlooked the presence of the species.

Certain categories of records appear underreported. Despite guarantees of anonymity, some members of the public were reluctant to provide reports of incidents involving the deliberate or accidental killing of a BFC for fear of prosecution or condemnation.

The incident rate of roadkill also appears to be low, although this is a subjective opinion of the researcher. Collinson (pers. comm., 2014), manager of the

Endangered Wildlife Trust’s Roadkill Research and Mitigation Project also reported that no records for the species had been received from the members of the public contributing roadkill data to the project in the period of 2013 – 2014

following its inception. The reasons for this could vary from simple misidentification by the public possibly linked to a lack of awareness of the species; the natural vigilance of the species, or to extremely low densities of cats and/or vehicular traffic in some parts of the country.

4.5.1.3 Research effort

Differing research efforts resulted in spatial and temporal biases and this was particularly apparent between the current record collecting period and the historical period. From the results, it could be inferred that BFC have virtually disappeared from the northern areas of its range, particularly in Namibia and

Botswana and currently only occurs in the central regions of South Africa. While it may be that South Africa and its central regions are, indeed, a stronghold for the species, this skewing of the data is more likely to be an artefact of research effort.

With the researcher being located centrally South Africa, it was logistically easier to establish and maintain public awareness in this area. In addition, a number of other research associates and professional problem animal control officers also made an effort to contribute data from specific areas in which they were conducting field work which resulted in most of the OR 3 ratings.

Increased research effort and interest in the species by the researcher and

BFCWG colleagues explains the sudden increase surge in temporal observations.

These increased significantly from 1990 onwards. Again, this should not be interpreted as an increase in population size but rather as the result of focussed field and literature investigations.

4.5.2 Range extensions

According to Rondinini and Boitani (2012:32), in order to determine the usefulness of a map, the following must be known: the method used to make the map; the data used; if the map is based on expert knowledge and the experts’ credentials; and, if the data is collected as point occurrences, how it was collected, the biological significance, the time span and how it was extrapolated. Doing this places the map in the appropriate context, acknowledges its limitations and ensures it is used appropriately.

Depending on the species, and the data and methods used to make them, a species’ distribution can be mapped in two ways: either estimating the EOO or the actual area occupied (AOO). All the maps in this section were calculated using

EOO, and is the method used in the IUCN Red List assessments when measuring geographical range sizes and determining extinction risks. According to Gaston and Fuller (2009:7) the EOO explicitly measures the overall geographic spread of the localities in which a species is found, and not the area over which it actually occurs. These maps are generally similar to those that appear in field guides.

Extent of occurrence maps differ to those depicting AOO, which simply put, counts the predetermined grid cells in which a species is present in a specific area or region. These maps tend to correlate much better with population size than EOO

(Gaston & Fuller, 2009:5).

In order to determine the distribution of the BFC, the authors all used the marginal occurrences mapping methods (Gaston & Fuller, 2009:2) which presumes the

outermost geographic locations to be the species’ range limit, and interpolating between these points to delineate the boundary. This is the most accurate method of measuring EOO although early maps were undoubtedly approximate with low ratios of genuine observational data to interpolate. Maps in modern field guides now incorporate additional information such as environmental conditions and seasonal information using computerised mapping techniques (Gaston & Fuller,

2009:4) which increases the overall accuracy.

However, Rondinini and Boitani (2012:32) warned of two common errors that readily occur in carnivore maps. These maps either erroneously indicate that a species is falsely present (error of commission) or falsely absent (errors of omission). Extent of occurrence maps are prone to commission errors as they intentionally ignore the internal extant of the area actually occupied by a species.

The historical maps presented here had EOO ranging from 1 236 178 km2 to

2 043 713 km2 in extent with a gradual linear increase in area size from 1982 to

2007. However, the first four maps from 1982 to 2005 all had locality omissions ranging from 17.6 to 26%, whilst the most recent (2007) only had a 4.1% error of omission.

Earlier mapping errors aside, there may also be other reasons that earlier geographical distribution maps had a higher rate of omission errors. Historical inaccuracies may have been unwittingly adopted from researcher to researcher that perpetuated earlier omissions due to further insufficient ground-truthing. Field investigations may have excluded some of the localities of earlier specimen

records due to the species no longer being present at that locality (e.g. a record near to Cape Town). However, the exclusion of newer data e.g. the Joubert,

Morsbach and Wallis (1982) report on the distribution patterns and status of some mammals in Namibia for the technical reasons explained earlier, constituted significant errors of omission. However, omissions may also have occurred due to scepticism on the part of the researchers who likely took into account the difficulty in validating observations due to the secretive, cryptic and naturally rareness of the species.

On the other hand, obtaining information for the creation of the new proposed map was greatly assisted by a few new technological advances only now available. A lot of information was obtained via public-sourced information aided by Internet- based social media platforms. Digitised museum collections and digitised literature are also now available on the Internet. This literature now also includes previously unpublished governmental reports (grey literature), proceedings of conferences, etc. All this new information, together with Geographic Information Systems (GIS) which allowed for mapping and statistical analysis, makes it likely that current proposed map (Figure 4.23) may be more inclusive and have less omission errors.

In the mapping process, an attempt was made to include potentially important records even if they were OR 1 such as the two outliers in the extreme southern region of Angola and those previously omitted from Namibia. The Angolan records are considered reliable (based on the reputability of the observers, available and suitable habitat features etc.) but were problematic in that the localities provided could not be precisely calculated to a specific QDS. For example, “Iona National

Park” was the western locality but it is a vast area that covers more than one QDS.

Other records received valid OR 1 ratings for being unreliable and extremely doubtful e.g. the Zambian record and were omitted in the mapping process.

However, the proposed northern map boundary is severely weakened by the lack of sampling localities in this region and as a result, there is a likelihood that an error of commission has occurred (Gastron & Fuller, 2009:3). Unfortunately, this uncertainty is a result of the political instability in Angola that does not allow for an inventory of its biodiversity (Ministry of Environment, 2009:24). However, if other factors such as habitat and human activities are taken into consideration Iona

National Park, for instance, does have suitable habitat in the form of discontinued coastal steppes consisting of sub-desert-like shrub vegetation dominated by

Aristida, Cissus, Salvadora and Welwitschia species. The park also houses a large human population of nomads whose main activity is stock-raising (Ministry of

Environment, 2009:24) which does not necessarily pose a threat to BFC.

Nevertheless, even if the species is reliably present in the northern reaches of the proposed EOO, Shortridge (1934: 96) said that even in 1930s skins were seldom brought in by natives for trading in Namibia by the Herero, Ovambo and Okavango tribes. Furthermore, the fact that they did not have a name for BFC may even be an indication of just how rare the species is in this region.

Very real range extensions were obvious to the eastern edge of the new proposed range, particularly in the Mpumalanga and KwaZulu-Natal area. Since there has not been wide-scale habitat change between the historical and current times, the

assumption is that the species was present in those areas previously, and just not documented. Again, the role of social media and the distribution of posters played a significant role in the creation of awareness of the species and the ongoing project in these areas. Nevertheless, and predictably, the marginal edges in the outermost limits had poorer sampling that confounds the depiction of an exact range edge.

Current BFC geographical distribution still suggests that the species remains absent from Lesotho and Swaziland. This may be because the species is absent from areas above 2 000 m (Sliwa, 2013:204), as are other potentially important sympatric species like mounding-building termite species (Ferrar, 1982:15) and springhares (Butynski, 2013:620).

Despite the range extensions, the mapping suggests that the current known BFC range does not suggest a disjunct distribution pattern, and that the species still has the most restricted range of all of the African cat species as indicated by Nowell and Jackson (1996:8).

It should also be remembered that EOO is a measure of range size and not an estimation of population size and as such, an extended geographical distribution does not necessary imply an increase in population size.

4.6 SUMMARY

The true range of a species is impossible to map particularly when the species is a small and rare carnivore like the BFC. Furthermore, distribution patterns are dynamic and can be expected to change over time in response to the species’ intrinsic and extrinsic variables.

Having established the distribution of BFC, it was possible to measure its range in terms of its EOO. It would appear that the historical geographical distributions from

1982-2007 have gradually increased the geographical distribution of the BFC but still underestimated the EOO of the species by both errors of omission and commission.

Since then, the map of Wilson and Sliwa (2007) appears to narrow the error of omission significantly and increased the range significantly. The current proposed map here has only omitted doubtful locality records and as a result has no errors of omission, and once again increased the predicted range of the species.

However, using this mapping technique now risks increased and unintentional errors of commission typical of EOO maps.

These range extensions over time cannot be inferred as true increases in the geographical distribution of the species, but rather as an artefact of possible under-reporting in the past.

CHAPTER 5

RESULTS AND DISCUSSION: CONSERVATION STATUS

5.1 INTRODUCTION

The conservation status of a species indicates whether the species still exists and how likely and over what time period it might become extinct in the future. The

IUCN Red List classifies species into nine conservation groups based on criteria that include: the rate of decline; population size; area of geographic distribution; and the degree of population and distribution fragmentation (IUCN, 2001).

This investigation attempts to determine if an estimated population size for the

BFC can be made using the updated geographical distribution information.

Additional conservation issues that arose during the research processes are explored here with the view to identifying all potential conservation threats faced by the species and ranking these in order of importance.

5.2 ESTIMATION OF POPULATION SIZE

The process of ranking species vulnerability by the IUCN uses a number of criteria with the size of the remaining population and its geographic range being the two main underlying factors that influence population size and extinction risk (Nowell &

Jackson, 1996:2-6).

Using 790 historical and current BFC locality records, the most inclusive distribution map to date has been created. However, the marginal occurrence mapping method used to make these basic field guide-type maps are estimates of extent of occurrence (EOO) and are designed to measure the area in which the species may occur. It is important to understand that this method is not a measure of population size. However, an accurate estimation of EOO is an essential part of the conservation assessment process and is used in the process to determine population size.

Extent of occurrence is also the basis for determining area of occupancy (AOO).

Of the 3216 QDS that fall within the EOO, only about 340 (10.6%) had records

(historical or current), totalling an AOO of 232 900 km2. Of the occupied QDS, 291 of these occurred in South Africa, covering 14.3% of the total number of QDS within the borders.

A kernel density estimation (KDE) heatmap (Figure 5.1) was created using the basic techniques employed for the review of the BFC conservation status (Wilson,

Sliwa & Drouilly, in prep.) in the current revision process of The Red List of

Mammals of South Africa, Swaziland and Lesotho. However, this mapping exercise was extended to include the entire southern African subregion rather than just South Africa, Swaziland and Lesotho. It also included both historical and current data. Areas on the map in red are considered high-density occurrence areas, yellow areas medium-density areas, dark green areas low density and with the remaining light zone as estimated range.

FIGURE 5.1: A kernel density estimation heatmap of black-footed cat distribution patterns in southern Africa (Map credit: L. Roxburgh)

The total geographical distribution (EOO) was calculated during the mapping exercise and was estimated to be 2 214 276 km2. The estimated areas of high,

medium and low density regions, and population size both nationally for South

Africa and regionally (globally) can be seen in Table 5.1.

TABLE 5.1: Estimated areas of high, medium and low density regions, and population size both nationally and globally

Area (km2) / Adult population size

Area High Density Medium Density Low Density Total (0.02 (0.01 (0.007 individuals/km2) individuals/km2) individuals/km2)

South 2 624 /124 975 2 515 / 179 654 3194 / 456 293 8 333 Africa

Regional 2 624 / 124 975 2 515 / 179 654 4 567 / 652 482 9 707 & Global

Areas of higher density appear to be clustered in the central regions of South

Africa, dropping substantially in a northward direction, but this may be linked in part to a reporting bias. However, based on the mapping results the regional

(global) population estimates are 2 624 (0.02 individuals/km2) individuals in the high density areas; 2515 (0.01 individuals/km2) in the medium density area, and

4 567 (0.007 individuals/km2) in the low density areas. This estimates a total of

9 707 mature animals within the entire southern African region. Summing the three density zones within the boundaries of South Africa results in a total estimated population of 8 333 mature individuals.

Estimates of population sizes are most sensitive to the size of the high density clusters. A more conservative estimate would yield a lower population size, while a less conservative estimate would yield a larger population size. Even increasing the high density areas to cover 70% of the observations of BFC, would still only yield a global population estimate of less than 12 000 individuals.

5.3 CURRENT CONSERVATION STATUS

Black-footed cats are readily acknowledged as an uncommon to rare species

(Shortridge, 1934:96; Smithers, 1971:128; Visser, 1976:73; Stuart, 1982:8; Stuart

& Wilson, 1988:23; Nowell & Jackson, 1996:8; Sunquist & Sunquist, 2002:80;

Skinner & Chimimba, 2005:406; Sliwa, 2013:204). The IUCN Red List Categories and Criteria were designed for global taxon assessments (IUCN, 2001:6).

However, they are often applied at a regional or national level, and when this occurs it may be that the national or regional category is not the same as the global category.

5.3.1 Global and regional status

Since 2001 the IUCN Red Data global and regional classifications have listed BFC in the threatened category, Vulnerable according to the criteria C2a(i). This criterion classifies the species as having: a total effective population size of 10 000 or less mature breeding individuals; no subpopulation containing more than 1 000 individuals, and with numbers likely to be declining (Sliwa, 2008). In the most

recent IUCN Red List reassessment (Sliwa, et al., 2015), the global IUCN conservation status has remained unchanged.

The IUCN/SCC Cat Specialist Group has BFC ranked as Category 2 for global vulnerability but ranked higher as Category 1 for regional vulnerability (Nowell &

Jackson, 1996:7). They are also considered the most vulnerable of the sub-

Saharan cat species and needing first attention in terms of conservation and species management plans. This category system ranks a species according to their vulnerability to extinction and assigns a score to four criteria (habitat association; size of geographical range, body size, and active threats against the species).

The species has also been included on Appendix 1 of CITES as an endangered species since 1975. Trade in species under this appendix is permitted only in exceptional circumstances. Hunting of BFC is prohibited in Botswana and South

Africa, but is not afforded legal protection in Mozambique, Namibia and Zimbabwe

(Nowell & Jackson, 1996:9). The species has no status in Angola as the fauna of the region is not well known due to the political instability in the country not allowing for an inventory of the biodiversity (Ministry of Environment, 2009:24).

5.3.2 National status

Currently, the species is listed nationally in the Red Data Book of the Mammals of

South Africa (2004) only as Least Concern (Friedman & Daly, 2004:172-173), a status it has held since 2002. The more recent conservation assessment carried

out in 2015 by Wilson, Sliwa & Drouilly (in prep.) for the revised version of The

Red List of Mammals of South Africa, Swaziland and Lesotho has seen the status uplifted to Vulnerable on the basis of the data presented in this thesis.

In South Africa BFC are also listed as a Protected Species under the Biodiversity

Act 10 of 2004 (SA, 2004) and were still included in the 2007 updated species list.

In addition, the species is considered a Near Endemic and to have a self- sustaining wilderness status which is currently unmanaged (Wilson, Sliwa &

Drouilly, in prep.).

5.3.3 Principal conservation threats

According to Nowell and Jackson (1996:7), Friedman and Daly (2004:172) and

Sliwa (2008), the currently acknowledged threats include:

 Human induced habitat loss as a result of overgrazing that leads to bush

encroachment and a loss of prey base due to the degradation processes.

 Indiscriminate persecution resulting in direct losses during predator control

efforts, as well as accidental poisonings during carcass baiting and locust

spraying activities.

 Has a highly restricted range.

 Low population size of less than 10 000 mature individuals remaining

throughout its entire range.

5.4 EMERGING ISSUES

A number of key issues were noted whilst reviewing literature and during the course of in-depth interviews during the geographic distribution survey process.

Many of these issues, as well a few additional risks, were discovered during the research investigations by the BFCWG underway since 2007 (Sliwa et al., 2007;

Sliwa et al., 2008; Sliwa et al., 2009a; Sliwa et al., 2009b; Sliwa et al., 2010; Sliwa et al., 2011; Sliwa et al., 2013; Sliwa et al., 2014; Sliwa et al., 2015). These issues have both natural and anthropologic factors involved and many may have significant conservation implications, either positive or negative, for the species.

5.4.1 Road mortalities

The extent of road mortality and the effect on local cat subpopulations is unknown.

The incident rate recorded in the Endangered Wildlife Trust’s Roadkill Research and Mitigation Project database indicated zero records up until December 2014 despite the project being underway since 2012 (Collinson, pers. comm., 2014).

Only 2.9% (n = 17) roadkill records were correctly reported (Figure 5.2) to, or directly observed (and collected as museum specimens) by, the researcher.

FIGURE 5.2: A BFC roadkill victim near Aberdeen submitted as a sighting record by a citizen scientist

Black-footed cats were roadkill victims on tar roads as well as on gravel roads.

However, there were an additional 22 (3.8%) records of responses in which people reported having seen BFC roadkill in the past (exact location and/or dates uncertain). Included in this calculation were records categorised as museum specimens or taxidermy mounts for which the origin was a collected roadkill and mounted as a trophy. In addition, a further 29 (5%) of the sight records came from

BFC observed on or crossing a road at night.

These additional records suggest the possibility that roadkill mortalities are underreported and that a much larger number of BFC die on public roads than can be determined conclusively at this time.

5.4.2 Hybridisation

Throughout the survey period, there were allegations of possible hybridisations of

BFC and domestic cats in several public responses and even in literature

(Shortridge, 1934:96; Rautenbach, 1982:150). These records were all discounted during the validation process. One 1945 museum record from the Karoo housed at the Amatole Museum, King Williamstown is accessioned as a hybrid with a domestic cat. During research on behalf of Peters, Baum, Peters and Tonkin-

Leyhausen (2007:221-337), more than 30 cranial measurements were taken of every skull from all the South African museums. This cat’s skull’s morphological measurements fell well within range of BFC of the same sex and age. A hybrid specimen would be expected to show different characteristics and it is therefore highly unlikely that it was a hybrid. No observations of hybrid individuals in the wild have been made either by the researcher or by Sliwa (pers. comm.) since 1992.

To date, the only confirmed hybrids were animals in a captive setting (Leyhausen,

1979:95).

5.4.3 Population genetics

Black-footed cats belong to the lineage of Old World domestic cats that emerged about 6.2 million years ago (Werdelin et al., 2010:61). The typical evolutionary lifespan of a mammal is about 1-10 million years from origination to extinction

(Mills, 2007:13) but this varies widely between taxa. Kuhn et al. (2011:7) recovered a fossil record of 1.977 million years old from the Malapa Hominin site

near Krugersdorp, Gauteng Province, which at that time was open woodland habitat.

This would indicate that as a species, BFC are genetically neither a new nor young species. However, Davis (S.a.) indicated that wild populations contain healthy levels of genetic diversity comparable to populations of other wildcat species.

Although there appeared little inbreeding in the wild BFC populations tested by the

BFCWG, there was some evidence of a possible bottleneck at some point. A population being researched near Kimberley, South Africa, has a unique genetic variant that suggested it might be a unique and important population.

Throughout the literature, two subspecies have been described and these persist in theory (Meester, et al., 1986:130-131). Felis nigripes nigripes, the nominate genotype form described by Burchell from the northern part of the range and Felis nigripes thomasi Shortridge, 1931 (Shortridge, 1931:119) from the southern and eastern parts of the range. The subspecies status has yet to be confirmed or discounted.

5.4.4 Fragmented distribution

The records and reporting rates in this study suggest that the central South African subregion remains the stronghold for the species with approximately half (50.4%) of records coming from this region. Whilst the geographical distribution or extent of occurrence (EOO) of the species may be greater than previously documented by other researchers, the area of occurrence (AOO) of the species, where it is known

and has been surveyed by the BFCWG (Sliwa et al., 2007; Sliwa et al., 2008;

Sliwa et al., 2009a; Sliwa et al., 2009b; Sliwa et al., 2010; Sliwa et al., 2011; Sliwa et al., 2013; Sliwa et al., 2014; Sliwa et al., 2015) is highly fragmented and patchy within the EOO. This assumption was supported during the mapping and estimations of population size during this research. It is expected that this effect is greater near the distributional boundaries, particularly in the north where the extreme environmental conditions like droughts and naturally low prey densities are even more challenging for the cats. Already, Keith (2005:193) has noted decreasing changes in the area and habitat quality of one of the key prey item species of BFC, the gerbil mouse (Malacothrix typica).

5.4.5 Habitat preferences

As indicated previously, all researchers agreed that the species is endemic to the arid grasslands, dwarf shrub, and savanna of the Karoo and Kalahari in southern

Africa, with a particular preference for panveld edge vegetation. Densities of BFC are expected to be higher in these habitats, and markedly lower in low-quality habitat (Sliwa, 2013:204).

Due to their elusive nature, BFC are generally known to avoid areas with significant human activity. However, during the course of this investigation, seven separate reports (0.9%) received were of BFC frequently seen in recently ploughed and/or planted agricultural lands. The cats were easily observed hunting, and it is likely that they are utilising the seasonal fluctuations of southern multimammate mouse (Mastomys coucha) and the Natal multimammate mouse

(M. natalensis). These pioneer rodent species are well-known for population explosions in agricultural areas where they are considered a pest, as well as degraded savannah veld and following fires (Avenant & Cavallini, 2007:217; Leirs,

Stenseth, Nichols, Hines, Verhagen et al., 2013:469).

5.4.6 Indiscriminate persecution

Black-footed cats have the potential to be by-catch victims of accidental poisonings (e.g. locust spraying, predator control lures/baits), as well as sport hunting throughout most of their range in South Africa (Nowell & Jackson, 1996;

Sliwa, 2008). There were a number of direct losses of individuals for these reasons.

There were 15 reports (1.8%) from interviewed farmers and problem control agents that admitted to either deliberately or accidentally hunting BFC, many claiming to have shot or trapped cats “several” times. A number of farmers steadfastly claimed that BFC are a problem-causing predator capable of hunting and killing young lambs and springbok, and that hunting them was justified. These were two separate reports of BFC raiding chicken coups resulting in the cats being trapped and killed (Figure 5.3).

FIGURE 5.3: A BFC on roadside display in the Karoo after being trapped and killed after raiding a chicken coup

However, most interviewees claimed to have accidentally shot BFC after mistaking the large eyes for that of a larger predator such as black-backed jackal or caracal, particularly at a distance. They appeared regretful for the mistake. During this period, there were no documented wide scale locust spraying activities in the region, and thus no reports of related BFC mortalities.

5.4.7 Industry interest and use

Surprisingly, there appeared an interest for this species in the trophy industry as witnessed by permit applications and requests for trophy mounts from local and international clients made to taxidermists (Wilson, Sliwa & Drouilly, in prep.). This happening, in spite of the fact that trade is not permitted for the species due to its

CITES I listing, and it being a Protected Species under the Biodiversity Act 10 of

2004 (SA, 2004) in South Africa.

Despite this, 16 (2% of the database records) taxidermy businesses in South

Africa admitted to having prepared taxidermy mounts for clients, of which only one indicated possessing a permit to do so. Another four records received were for animals deliberately hunted as trophy animals in the Eastern Cape and Northern

Cape provinces. Two of these became museum specimens, one resulted in prosecution, and the remaining trophy’s outcome unknown. Only a small proportion of the approximately 500 taxidermy practitioners in South Africa were approached in this investigation. It is also doubtful that not all the taxidermists were truthful in their responses to the survey questions for fear of persecution.

It was more difficult to assess the impact of traditional use has on BFC in southern

Africa. In Namibia during the 1930s Shortridge (1934:96) attempted to make a karros of skins, but only obtained 15 skins in a 12-year period because the skins were so seldom brought in by natives for trading. The Herero, Ovambo and

Okavango tribes did not have a name for BFC and referred to as “Sebala” meaning rare. This was probably a measure of how seldom they encountered the species.

Sunquist and Sunquist (2002:80) reported that a skin buyer in South Africa bought thousands of small animals in 1967, but only 19 of those were BFC. During this current investigation, there was only one record of traditional use. Amusingly, in

2007, Professor Anne Rasa, owner of Kalahari Trails Nature Reserve at the

southern end of the Kalahari Desert in South Africa, reported a “Bushman wearing a black-footed cat on his bum”. On questioning, he reported to her that he had picked up the cat as a roadkill 5 km from Andriesvale and that he considered them rare in the area.

Since the inception of the BFCWG and the subsequent field surveys; formal and informal presentations; this research carried out for this project, and regular media items, there has been a growing public awareness of the species. As a result, the

BFCWG receives regular enquiries to host scientific safaris (Figure 5.4) to view radio-collared individuals at its research site on the De Beers-owned Benfontein

Game Farm35 near to Kimberley, Northern Cape. The hosting of scientific tourists is now a regular occurrence, with many groups donating funds to the Group towards the running cost of the project. A nearby privately owned game farm,

Marrick Safari36, also regularly hosts safari groups specifically hoping to catch a glimpse of a resident, one-eyed, male BFC that is often seen on a small pan on the property.

35 www.diamondroute.com/explore-benfontein.htm

36 www.marricksafari.com

FIGURE 5.4: The researcher hosts scientific safaris for numerous groups and local television news crews in a bid to raise awareness about the black-footed cat

5.4.8 Reliance on sympatric species

In the wild, BFC are unable to create or maintain their own dens or burrows. Over time, BFC have become dependent (inquilinous) on abandoned or hollowed out termite mounds particularly when females have kittens (Figure 5.5), or the burrows of other small animals such as springhares and Cape ground squirrels.

During the investigation, the reliance on springhares only became apparent later in the field investigations by the BFCWG and the researcher. When interviewed, 20 respondents reported regular sightings of cats in areas where springhares were abundant. Unfortunately, this information was not available in literature surveys or with reported sightings via other sources such as MammalMap. This inquilinous association appeared more vital particularly when termite mounds became few or absent. In areas surveyed, the absence of both termite mounds and springhares usually appeared to result in the complete absence of BFC.

FIGURE 5.5: A black-footed cat kitten in a hollowed out snouted harvester termite (Trinervitermes trinervoides) mound (Photograph: A. Sliwa)

The intersect between the geographical range of springhares (Butynski, 2013:620;

MammalMAP, 2015) and BFC is indicated in Figure 5.6. This area of intersect is

1 938 682 km2 in extent meaning that at least 87.6% of the BFC range occurs in areas where springhare occur.

FIGURE 5.6: Black-footed cat geographical distribution (blue) overlying the range of springhare (grey) indicating am 87.6% intersection with BFC range

5.4.9 Intraguild predation

As a small species in a largely tree-less environment, BFC are vulnerable to larger predators like black-backed jackals, caracals and large owl species. This often leads to natural intraguild predation, namely the killing and eating of supposedly competing species that use similar resources are thus considered potential competitors for the larger species. Sliwa, Wilson and Kamler all personally witnessed interactions between black-backed jackals and caracals, none of which ended in a fatality. However, annually the BFCWG loses a varying number of all radio-collared project cats, many of these loses suspected (sometimes up to 50%) to be caused by larger predators such as black-backed jackals and caracal (e.g.

Sliwa et al., 2007; Sliwa et al., 2011).

Typically, mortalities caused by caracal resulted in significant signs of “overkill” usually consisting of multiple bite wounds and broken bones to the BFC, but the carcass remaining uneaten and the radio collar equipment undamaged and intact

(Figure 5.7 and Figure 5.8). However, mortalities caused by black-backed jackals normally resulted in the cat being devoured and recovered radio collars usually having significant chewing damage too (Figure 5.9).

FIGURE 5.7: The discarded and uneaten carcass of “Maya” a young female black- footed cat killed by a caracal in 2007

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FIGURE 5.8: “Maya”, a young female black-footed cat killed by a caracal in 2007 showing bite wounds circled in red

FIGURE 5.9: The only remains of “Panga”, a large radio-collared male black- footed cat that was killed and eaten by black-backed jackals in 2007

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Whilst intraguild predation in the wild can be considered a natural occurrence, the

BFCWG also has evidence of BFC having been killed by traditional herding and hunting dogs (Sliwa et al., 2013). In addition, 10 (1.3%) records in the database were reports of interactions with BFC and dogs, with nine resulting in fatalities

(sometimes involving more than one BFC dying during the attack, e.g. entire kitten litter, or female and kittens), and only one instance of the cat actually escaping in the attack. The situations varied with the attacks occurring in the veld to taking place in or near farmyards and homesteads.

In the Eastern Cape, and occasionally elsewhere in the country, Anatolian shepherd dogs are popular breeds used to guard livestock against jackals, servals, caracals and leopards. A number of farmers expressed concern during the study that these dogs had completely eradicated all of the small mammals on their farms, and surrounding areas on occasions when they wandered off. The victims included non-target species such as hares, springhares, mongooses, aardwolves (Proteles cristatus), bat-eared foxes (Otocyon megalotis), aardvark and small antelope species, as well as livestock.

5.4.10 Natural catastrophes

It would seem that BFC are vulnerable to natural disasters, many of which could potentially cause mortalities. During the study period, the BFCWG observed at least two such mortalities (e.g. Sliwa et al., 2009), although several other weather- related deaths of radio-collared individuals were suspected.

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One female BFC died after being caught aboveground in a severe storm (Figure

5.10) and another died after a den was flooded and collapsed (Figure 5.11) with the cat still inside.

FIGURE 5.10: The remains of “Gogo”, a radio-collared female black-footed cat that died aboveground, possible due to a severe storm in 2009

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FIGURE 5.11: The remains of “Jimbo”, a radio-collared male black-footed cat that died in a den that collapsed after it was flooded in 2009 (Photograph: A. Sliwa)

5.4.11 Diseases

Black-footed cats share their territory, prey base, parasites as well as infectious disease susceptibility with many small carnivores including genets, caracals,

African wildcats, yellow mongooses (Cynictis penicillata), suricates (Suricata suricatta), Cape foxes (Vulpes chama), bat-eared foxes, black-backed jackals, aardwolves, striped polecats (Ictonyx striatus), and even domestic dogs and cats.

This provides numerous opportunities for disease transmission (Lamberski et al.,

2009). According to Lamberski et al., (2009) preliminary results from 23 BFC tested near Kimberley and De Aar, Northern Cape, South Africa, suggest that 25% of the BFC tested were seropositive for canine distemper, whilst 30% were positive for feline calici virus and 13% for West Nile virus. At that time, there was

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no serological evidence for feline parvovirus, feline herpes virus, feline calicivirus, feline immunodeficiency virus, toxoplasmosis or rabies.

In captivity, BFC develop respiratory diseases; are susceptible to toxoplasmosis, and show a high prevalence for AA-amyloidosis. The latter is a disease characterised by fibrillar protein depositions in many organs usually cumulating in renal failure. About 70% of the documented deaths of captive cats internationally are because of this disease and although initially thought that this disease was a captivity-induced disease, the presence of amyloid in a free-ranging black-footed cat was detected by Terio et al. (2008) and Zimmermann Lawrenz and Sliwa

(2011). This provides additional evidence for a species predilection and supports the existence of a possible familial type of amyloidosis in BFC. Habitat fragmentation and population isolation can only exacerbate this disease at subpopulation levels, whilst at the same time, the disease is the major reason why currently global captive breeding programmes are not self-sustainable (Sliwa, pers. comm., December 2014).

5.4.12 Climate change

Since the Pleistocene period, it is highly probable that the range of the BFC has shrunk southwards as the nature of the environment in which the cats previously lived has transformed because of natural climate changes (Kuhn, pers. comm.,

2013). Whilst actual climate change over the period of this study cannot be measured or inferred directly to BFC, certain aspects can be examined. Fordham,

Akcakaya, Brook, Rodriguez, Alves et al. (2013:901) have predicted that the

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impact of rising temperatures and changing rainfall patterns on the prey of the

Iberian lynx may lead to its extinction in as little as 50 years unless there is a dramatic shift in the conservation strategies for the species. It is suggested that anthropogenic factors resulting in habitat fragmentation and changes in the prey base may be worsened by the effects of global warming caused by increasing greenhouse-gas emissions.

Nkunika, Shiday, Sileshi, French, Nyeko et al., (2010) has reported that climate change will affect termite distributions in Africa. According to these researchers, the areas most likely to be affected in the southern African subregion will be north- eastern Namibia, all of Botswana except the southern most areas, and the western and south-western parts of Zimbabwe. This may be critical habitat to BFC, especially in Botswana. Here, springhares are already under pressure (Butynski,

2013:623) and this increases the need for available termite mounds. Fortunately, termite species are adapted to hot and arid environments are actually predicted to expand their range.

5.4.13 Presence in formally protected areas

Of the 790 records, there was a relatively poor reporting rate from formally protected areas (FPA) throughout the region. These are areas protected either by national legislation or by provincial ordinances and are actively administered in accordance with management plans. These areas, deemed national assets, are unlikely to be sold or undergo a change in land-use in the near future. Since these areas are considered to be safeguarded from anthropogenic influences and thus

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relatively stable over time, both historical and current records were analysed collectively.

The total number of FPA for each country reported to have BFC present can be seen in Table 5.2, together with the number of FPA in which BFC have been recorded. The regional Red Data assessment currently underway (Wilson, Sliwa &

Drouilly, in prep.) allows for a 50 km buffer zone radius around each locality record. This is deemed the maximum dispersal capability of an individual. Twelve additional FPA’s fell within these buffer zones and were added to the table because the possibility exists that BFC could disperse to these areas or may already be present but are, as yet, undetected.

TABLE 5.2: Formally protected areas in southern Africa indicating the presence/absence of black-footed cat within the boundaries, or within a 50 km radius of the nearest black-footed cat locality record

Number of Number of FPA Total Number of FPA with within a 50 km Country number of FPA within BFC radius of a BFC FPA the BFC EOO records record South Africa 98 49 11 10

Botswana 11 11 4 0

Namibia 14 7 1 2

Angola 14 2 1 0

Zimbabwe 44 1 1 0

Total 181 70 17 12

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Of the 181 formally protected areas in the region, 70 (40%) fell within the BFC geographical distribution or EOO. The highest representation (100%) was in

Botswana and the lowest in Zimbabwe with only one of the 44 FPA present there having reported BFC.

There were 24 locality records from 11 different FPA in South Africa, but BFC were only regularly reported from three (3.1%) namely Karoo National Park,

Mountain Zebra National Park and SA Lombard Nature Reserve. All other FPA records were single observations and no further reports of the species were made to the researcher. Additional, but unconfirmed records were reported from Addo

Elephant National Park but these need further investigation.

All of the Botswana BFC records occurred in four (36.4%) of its FPA, with multiple records in all four areas, namely Kgalagadi Transfrontier Park (historically),

Central Kalahari Game Reserve, Makgadikgadi Pans National Park and Moremi

Game Reserve. Of the 14 protected areas in Namibia, only one BFC record was located within the boundaries of a FPA, namely Bwabwata National Park in north- eastern Namibia. There were a number of records near Etosha National Park as well as Waterberg Plateau Park. However, due to the clumping of 309 records into just 12 district localities because of insufficient locality details from the Joubert,

Morsbach and Wallis (1982) survey report, other records may have fallen within 50 km of a FPA too, but could not be assessed here.

There was only one FPA record from Angola, namely Iona National Park, in the extreme southwest. The sole Zimbabwean record to date was from the western

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edge of Hwange National Park on the Botswana-Zimbabwe border. Records into these countries are considered only as marginal range extensions at this time, pending further information forthcoming from these regions.

5.5 DISCUSSION

A variety of both intrinsic traits and extrinsic factors may be placing this species at a higher risk of expedited population declines. These issues can be either natural or anthropogenic (or a combination of both) in nature. According to Purvis,

Gittleman, Cowlishaw and Mace (2000:1947), intrinsic factors in carnivores include being at a high trophic level; low population density; slow life history, and small geographical range size. These traits are significantly and independently associated with a high extinction risk in declining species and account for 50% of the driving forces behind the extinction risk. The remaining causes of extinction are usually due to external anthropogenic factors, which affect not only a specific species, but also a whole host of organisms in the same environment.

5.5.1 Range and population size

Previous inaccuracies in population size estimations have BFC incorrectly listed as

Least Concern in South Africa, where the largest proportion of its range occurs.

These estimates were confounded by a lack of consistent research and reporting efforts. It was necessary to incorporate both historical and current BFC locality records in the population size mapping exercise due to the lack of temporal and spatial data from Namibia and Botswana. This then erroneously assumes that

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historical populations are still extant and unchanged spatially or temporarily, which is probably unrealistic. The results are also confounded by reporting biases in

South Africa.

However, perhaps more important than an exact population size estimation, the heatmapping exercise offers a replicable method that can be used within the assessment region that allows for future spatial and temporal comparisons.

The kernel density estimation (KDE) map graphically suggests the AOO to be significantly smaller than the EOO, even when records are buffered. Fine-scale resolutions of 2 km are normally used by the IUCN during conservation assessments. However, since individuals have the potential to disperse up to 50 km, it was decided to use a coarser grid of standard QDS to determine presence or absence within the EOO. At roughly 27.4 km north-south and 25 km east-west

(totalling around 685 km2), a QDS is significantly smaller than the estimated dispersal area of a circle with a 50 km radius that produces an area of about 8 000 km2 in size. As such, there are records of BFC from only 10.6% of the EOO. Most species usually only occupy 40-70% of their EOO (Jetz, Sekercioglu & Watson,

2008:110) and species that occupy less are considered at-risk species and more likely to have range overestimations. This KDE mapping method also still assumes that even in the low-density areas, cats were present, albeit in very low numbers, even though it is with some certainty that the species does not occur in some areas for example, Bushmanland (Wilson, pers. obs.) or the Kgalagadi

Transfrontier Park currently (Skinner & Chimimba, 2005:407).

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It is also not possible to determine the extent or number of subpopulations for

BFC, but it appears that populations are severely fragmented and isolated (Wilson,

Sliwa & Drouilly, in prep.). It is certainly unlikely that any cluster numbers more than 1 000 individuals, which is one of the conservation measures used to assess declining populations. The current newly proposed uplifting of the South African conservation status from Least Concern to Vulnerable is fully justified as is the continued global status of Vulnerable since both populations are estimated to be below 10 000 mature cats, either individually or collectively.

The strengths of the KDE map should not only be seen in terms of population estimates, but also in terms of spatial density of the species across its range. Only the South African population size can be reliably assessed at this time due to a good sample size of current geographical localities. Once again, the central regions of South Africa indicated overall higher population densities, but for the first time, other regions were also indicated in the Eastern Cape and Western

Cape, as well as in the far eastern range extension towards northern KwaZulu-

Natal. Even North West province, an area historically excluded from distribution maps now suggests viable populations of BFC in the province. Densities appear much lower on the extremities and in the northern areas of the geographical range of the species, and it probable that this is accurate, even had there been more locality records from these areas.

Given the strong indication that this is an at-risk species, further investigations into the known and emerging issues were deemed necessary in this study.

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5.5.2 Anthropogenic threats

Human-initiated habitat destruction is currently one of the leading anthropogenic causes of species extinctions. It alters the habitat in various ways, either directly or indirectly, to such an extent that a species is no longer able to survive or thrive. A number of risks were identified and quantified or explained.

Habitat fragmentation results in the creation of island populations resulting in limited dispersal opportunities and potentially restricting genetic exchange between subpopulations. As wild populations become genetically isolated, reduced genetic variability threatens population viability by increasing susceptibility to disease and reducing reproductive fitness and the affected species may not recover from these impacts (McManus, Dalton, Kotzé, Smuts, Dickman et al.,

2015:336).

Habitat loss or changes have already been considered the most severe threat to

BFC in previous conservation assessments. Habitat degradation that results in the loss of prey base is a serious threat, but an additional consequence of habitat fragmentation and population isolation is increased contact with other carnivores and the pathogens that they carry that could potentially be detrimental to entire local populations.

Whilst roads do not constitute a serious cause of habitat fragmentation for BFC, the effects of roads on wildlife can be significant. However, understanding the causes and patterns is confounded by the complexities of differing habitats,

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dynamics of animal populations and their behaviour, as well as the anthropogenic factors such as road design, vehicle types and driver attitudes.

Black-footed cats range widely throughout their habitat and a tar or gravel road does not constitute a barrier for the species. Having a small body size means it does not pose a real threat to any forms of vehicular traffic and most drivers would not be forced to actively avoid striking cats on a road. Once dead, the tiny body will rapidly disintegrate with additional crushing by other vehicular traffic.

Carcasses that remain intact are easily overlooked on the roadside or dragged out-of-sight by scavenging crows and jackals. Therefore, despite the relative alertness character of the BFC, it is likely to be roadkill victim, and one that is frequently undercounted. In a region with an already vast and growing road network, this poses a growing and an unquantifiable risk.

Another thought is that road verges are more ecologically productive ecotones due to microclimatic changes and that they hold both higher densities and a wider variety of species (Coffin, 2007:402). This in turn makes them more attractive to some species, particularly those that use them as corridors or conduits (Coffin,

2007:400). Although BFC appear to use roads for moving about during the night infrequently (Sliwa, pers. comm., 2015) both tarred and dirt roads within their territories will need to be crossed from time to time. These roads and the associated harvested roadside verges may on occasion also offer easier movements than in adjacent, denser habitats for the short-legged cats.

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Southern Africa has undergone many habitat changes over the millennia. Two million years ago, fossil BFC living at the Malapa Hominin site discovered by Kuhn et al. (2011:7) would have lived in a markedly different habitat to that of today. During that period, the dominant vegetation in the region was cooler, moister, and of a forest vegetation type (Bamford, Neumann, Pereira, Scott, Dirks et al., 2010:27). Kuhn (pers. comm., 2013) speculated that this may have indicated that, as a non-tree climbing species, BFC with their short legs were a forest floor- dwelling species. Their spotted pelage would have blended in well with the sunlight-dappled, leafy undergrowth where they would have been hunting and escaping detection from larger predators.

Remarkably, BFC are still present in the region despite the changing environment and climate and have even managed to thrive in harsh semi-arid and arid environments. This suggests a degree of resilience and adaptability. Spatial- temporal partitioning is an important mechanism for minimising resource competition among sympatric species (Ramesh, Kalle, Sankar & Qureshi,

2012:269). How sympatric species with similar diets are able to coexist depends largely on several factors such as physiological adaptations, prey availability and hunting strategies. Having short legs and a small body size, may be morphologically advantaged to maximise the lack of ground cover in agricultural situations than, say, the larger African wildcats in the same situation. Thus, BFC are able to exploit the seasonal rodent explosions associated with sowing and harvesting activities. However, given the unpredictable nature of these events and associated risks due to the proximity to humans and their domestic carnivores, the benefit to resident BFC is uncertain.

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Despite a number of landowners and problem control agents admitting to deliberately or accidentally hunting and killing BFC, the suspicion is that a larger number of cats are killed than is reported on and confounding the statistics of this threat. Given that populations are slow to recover in these situations, the impact is potentially a very serious threat to localised populations.

Indiscriminate prosecution has already been a previously documented threat, and this appears unlikely to abate. The use of predator control methods such as cage traps, foothold (gin) traps, and poison baits is widespread throughout the region regardless of legislations governing the practices. A number of interviewed farmers admitted to resorting to these measures in desperation when facing rising stock losses attributed to predators. However, the reality behind the problem is that these conflict situations are often the result of changing and poor farming practices (Nattrass & Conradie, 2013:11). In many instances, the farmer is not personally responsible for setting and clearing these devices, or applying poison baits. This exercise is left to either a skilled farm employee or a compensated nomadic vermin hunter. Since non-target trap species such as polecats, mongooses and BFC are not paid bounty fees by the farmer, the carcases of these species are normally discarded in the veld and subsequently go unrecorded.

It is thus not possible to determine the full impact of these activities on the species.

The discovery of BFC trophy animals in the hunting fraternity was both surprising and disturbing. According to Van der Merwe, Saayman and Rossouw (2014:380), a survey carried out by the North West University in 2010 indicated that South

Africa’s hunting industry contributed nearly R6 billion to the economy for the 2009-

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2010 season. It makes sense that the safari hunting business has become increasing competitive. Safari operators booking clients are anxious to offer interesting and exceptional trophy species. This is one of the leading reasons why colour-variants such as golden gemsbok or black impala are currently so popular in the game breeding industry. Many safari operators even offer aardwolf, Cape fox, bat-eared fox, African civet (Civettictis civetta) and even aardvark (antbear) to clients37. Despite its small size, the BFC would make an excellent and challenging rare species to hunt, especially to an uncaring and ill-informed safari client that has hunted all the usual African predator species.

On a more positive note, BFC appear to have some appeal in the tourism industry due to a growing awareness of the species, in part, because of the work carried out by the BFCWG. Something of an iconic flagship species of the Karoo region, it is becoming a much sort-after species to see on nocturnal game drive safaris and a number of tour operators and wildlife preserves now offer this species as a tourism draw card.

Many of Africa’s wildlife species are extensively and uncontrollably utilised for traditional purposes. Whilst there are Bushman (Khoisan) legends about the less- than-imposing 20 cm high BFC being able to kill giraffe (Sunquist & Sunquist,

2002:78), this is more testimony to their ferocity rather than the actual capability to carry out this act. The rareness and inaccessibility of the species has prevented it

37 Example: www.selectsafaris.co.za/trophies/africagame.html (still valid 15 April

2015)

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from becoming a regular trade or medicinal item in any of the southern African indigenous groups and this situation is, fortunately, likely to remain so.

When species become threatened, often their continued existence is dependent on the formation of formal sanctuaries that protect them from anthropogenic influences. Examination of distribution data in this study clearly indicates that the majority of BFC records occur outside formally protected areas (FPA). The lack of presence in Zimbabwe and Angola in FPA is not of particular concern since the species appears only to have marginally range extension into these countries.

The presence/absence of BFC in FPA in SA needs to be determined with some urgency, especially since this region appears to contain the majority of the specie’s range and population. Gelderblom, Bronner, Lomard and Taylor (1995: 4) already acknowledged this 20 years ago, when they only recorded one of 74 BFC sightings in a protected area during a survey in the Eastern Cape. Even then,

Gelderblom et al. (1995:113-114) specifically stated that this species was one that required additional protection within formally protected areas as well as in biodiversity strategic plans.

The reasons for poor representation in FPA are varied. Medium- and larger-sized predator densities are often higher within the boundaries of FPA. These areas are often established to protect these very species from human-wildlife conflict, and in response to tourism demands to see larger carnivores. The intraguild risks for BFC are therefore higher within these areas meaning that they may actively avoid the area despite it having suitable habitat. Many protected areas are also too small to

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adequately conserve a full subpopulation of BFC with their large home range.

Other FPA also simply do not contain suitable habitat, e.g. mountainous areas above 2 000 m.

All the countries in southern Africa also have a large number of privately-owned and managed wildlife preserves and ranches of varying size. Many of them are likely to contain suitable habitat for BFC. However, it would seem that regions involved in stock farming with predator control measures in place (e.g. merino sheep farming in the Karoo) are likely to be more suitable BFC habitat. The implication of having a low representation of BFC within FPA is that the conservation of the species may rely on the cooperation of private landowners and managers.

5.5.3 Natural threats

Every species has a range of intrinsic threats, many of these acquired over time.

Hybridisation between domestic and wild animals is a major concern for biodiversity conservation, and as habitats become increasingly fragmented, conserving biodiversity at all levels, including genetic, becomes increasingly important. Both Shortridge (1934:96) and Rautenbach (1982:150) suggested possible hybridisation with domestic cats. Rautenbach further mentioned a highly unlikely example of a hybrid cat “seen in the business centre of Pretoria on a

Saturday morning”. However, Král & Zima (1980) in Wozencraft (1993:536) noted a distinctly different karotype from other Felis, and Lopez et al. (1997:281) reported distinct DNA sequences of this species that set it apart from the other

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Felis spp. concluding that the species evolved separately from an early point of the group’s establishment.

Fortunately, unlike the African wild cat (Le Roux, Foxcroft, Herbst & MacFadyen,

2015:288), this species does not hybridise easily with other cat species and has a very low risk of natural hybridisation under natural conditions. It is likely that so- called hybrid reports in literature were mistaken identifications of juvenile African wildcats, which share a remarkable resemblance with BFC until the intense spotting fades and the paler, uniform adult coloration develops. Genetically, wild

BFC currently appears to have healthy population genetics. However, with limited samples having been collected by the BFCWG and analysed to date, this status may change with broader spatial sampling.

According to the Hardy-Weinberg Principle, five conditions must be met in order for the genotype to remain constant in a species. These include the presence of a large population; random mating; no gene flow; no natural selection, and no mutations. McManus et al. (2014:338) have already shown that the solitary top carnivore African leopard populations can become genetically isolated within a few generations. The BFC too, is solitary and considered to have a restricted and fragmented distribution with no subpopulation containing more than 1 000 solitary individuals with limited dispersal capabilities (Wilson, Sliwa & Drouilly, in prep.).

This may increase the genetic risks for the species and prevent it from maintaining its genetic equilibrium.

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Genetic resolution needs to be obtained about the BFC subspecies, as well as further investigations into subpopulations to determine localised genetic diversity.

Distribution patterns do not indicate any geographical or ecological barriers that could have initiated genetic drift (Hoskin et al., 2005:1353) although BFC do show some geocline variations towards the extremes of its range (Sliwa, 2013:203). It is likely that the subspecies status is invalid.

However, the potential existence of two distinct subspecies, until ruled out, may have an impact on the approximately 75 BFC captive individuals worldwide. These cats are managed cooperatively as a single population by the Association of Zoos and Aquariums (AZA) Black-footed Cat (BFC) Species Survival Plan®38 (SSP) in

19 AZA-accredited zoos or accredited facilities in North America (Lamberski, S.a.).

The goal of the SSP is to create a sustainable population that is capable of maintain a 90% genetic diversity in the captive population for 100 years. This species is highly stressful by nature and notoriously difficult to keep in captivity.

Currently the captive population is only at 84% genetic diversity but is predicted to drop to 57% in the next century (Lamberski, S.a). Previously there were only a handful of cats in European zoos, with the last one dying in September 2014

(Sliwa, pers. comm., April 2015) and none in South African zoos. The introduction of new founder animals to improve the genetic diversity is therefore technically challenging. Sourcing from private collections is problematic because the origin and lineage of these individuals is not always known. This may result in a genetic

38 www.aza.org

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contamination between the two subspecies, should it be proven that they are valid subspecies.

This situation occurred in the 1970s and was discovered during the course of this investigation. A well-known and infamous South African zoologist and his colleague colluded to export BFC to zoos worldwide using permits issued in the

Eastern Cape for a number of F. n. thomasi. Unable to obtain the numbers of individuals listed on the permit, cats were sourced from the Northern Cape F. n. nigripes population. This was never declared to the legislative authorities or to the recipient zoos.

Fortunately, the future demands to improve genetic diversity in captivity may have a technical solution that will reduce the need to source wild-caught animals. As part of the work being carried out by the BFCWG, Herrick, Campbell, Levens,

Moore, Benson et al. (2010:552) assisted reproductive technologies such as artificial insemination, in vitro fertilisation and embryo transfers between captive animals are being developed. In addition, sperm from free-ranging male BFC is routinely collected during fieldwork and these frozen spermatozoa can be infused into captive females to further boost genetic variability in zoo populations.

The link between BFC and other sympatric species bears further investigation.

Similar to BFC, Skinner and Chimimba (2005:101) indicated that springhare distribution is patchy throughout its range. Both BFC (Sliwa, 2013:295) and springhares (Skinner & Chimimba, 2005:103; Butynksi, 2013:621) demonstrate a lack of dependence on water. Termites (Ferrar, 1982:41) and springhares

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(Butynski, 2013:620) do not occur above altitudes of 2 000 m, and neither do BFC

(Sliwa, 2013:204). Whether or not the absence of BFC from higher altitudes is linked to these species bears further investigation.

Whilst distribution maps do not exist for termite species in southern Africa, they are considered widespread with 165 species in the region (Nkunika et al. 2010). A number of termite species build mounds in this region, which are utilised by BFC particularly when females have kittens. Most commonly these species are

Trinervitermes trinervoides (in South Africa south of the Limpopo River) and T. rhodesiensis (north of the Limpopo in Botswana and Zimbabwe). Termite mounds are often destroyed by farmers during bouts of drought as the grass-harvesting activities by the termites are thought to compete with livestock grazing for resources.

Of particular concern noted in this research was the presence or absence of springhares in certain areas and regions. This in turn will affect the availability of burrows that BFC use for refuges. Unfortunately, springhares were often considered as a problem or damage-causing species requiring some control measures by farmers interviewed for this survey. However, more alarming is that, in Botswana, historical data indicates that 2.5 million springhares are harvested annually for the bushmeat industry (Butynski, 1973:208). With the practice ongoing, BOSTID (1991:194) reported that at least 3.3 million kilograms of springhare meat reached the market, and was the main source of bushmeat for human consumption in the country.

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Springhares are also often considered as a problem or damage-causing species requiring some control measures by farmers, with a number of farmers admitting to regularly hunting them because of this, but also for sport. Although a rodent, springhares are long-lived with a slow reproductive rate and they do not recover easily from a severe reduction in numbers. Together, the unregulated subsistence hunting of the springhares for bushmeat as well as its persecution as a problem species or for sport hunting may be leading to a possible eradication of the species in some regions, and with them, potentially the BFC in those areas. The overall implications of this are that changes in springhare density and range may well have an effect on BFC population sizes.

The association with springhares also raises an interesting possible range extension. The two species have strongly correlated geographical distributions ranges, although the springhare range extends much further north. This brings into question the possible validity of the Zambian BFC sighting. Since springhares are present in that area, is it possible that BFC occur there too?

Natural selection of BFC is very dependent on its ability to co-evolve with its prey and the environment. When a species is no longer able to evolve at the same rate as its environment, prey and predators, it is unlikely to survive and more likely to become extinct. It could be speculated that if mound-building termites were to somehow evolve and begin living in smaller groups, or have severe range shifts due to global warming; or the body shape and size of springhares were to alter,

BFC would have to evolve in order to continue a sympatric relationship.

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Should future investigations prove an important inquilinous relationship with springhares, the occurrence of springhares further north of the current known BFC range may suggest that the species could occur northwards as the isolated reports suggest. Given that BFC are harder to observe, occur at significantly lower densities, and have little economic value compared to that of springhares in these areas, it may account for the reason that this species having yet to be reported in these areas.

Intraguild predation was a newly documented risk for the species. Predation is clearly a high, natural risk for the smallest felid in the region. However, Sunquist and Sunquist (2002:80) suggested that BFC are adapted to this stress in several ways: females have a short estrus and receptivity period, followed by a long gestation period and small litter size, and the rapid development of the young reduces the time kittens are vulnerable.

However, in general, numbers of BFC and other larger carnivores are still negatively related (Bagniewska & Kamler, 2013:566). Kamler, Stenekewitz, Sliwa,

Wilson, Lamberski et al. (2015:125) stated that, despite partitioning resources differently among other sympatric canids that facilitates coexistence with the larger carnivore species, BFC home ranges still completely overlapped with those of the other carnivore species. This was supported and quantifiable to a certain extent by long-term field studies conducted by the BFCWG that suggested that intraguild predation is still one of the highest natural threats to the species.

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Ironically, the judicious management of black-backed jackals and caracal in the sheep-farming Karoo may have created a less competitive environment for BFC in which to exist. This may, in part, also be contributing to assumptions that the species stronghold in terms of habitat and population density may well be in this region.

However, the popularity of Anatolian shepherd dogs with sheep farmers is potentially a new emerging type of unnatural intraguild predation. These dogs are renowned for their success at removing all potential predator species near the stock they are guarding (Potgieter, Marker, Avenant & Kerley, 2014:404).

Unfortunately, many non-target species are also eradicated in the process. Whilst the cats are likely to avoid areas with Anatolians present, chance encounters could be fatal as the speed and digging skills of the dogs far outmatch those of this tiny cat.

In areas where hunting of predators using pack dogs (e.g. Eastern Cape), as well as traditional hunting with dogs (e.g. Zulus in KwaZulu-Natal) is routinely carried out, it is also unlikely that a BFC chanced upon could successfully escape. This would also apply to areas that have packs of roaming feral dogs too.

Many wild animals have to deal with seasonal climate disasters, as well as erratic occurrences of other natural disasters, although typically these are brief events.

The negative impact is usually brought on by the harm to their habitat rather than to the animals themselves. However, there are events from which they are unable to escape such as severe storms, flooding and fires. Natural disasters are

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responsible for killing, displacing and trapping animals in their refuges or even starving them when their prey base is destroyed. The ability of species such as

BFC to survive these events has not been determined, but providing the calamity is limited in extent and duration, repopulation from neighbouring areas will potentially occur over time.

The conservation and management of endangered carnivores provides limited opportunities to detect, monitor and contain diseases which are typically unpredictable. Repeated exposure of a species to diseases in their natural environment requires the species to develop a form of evolutionary resistance.

Whilst the solitary nature of BFC limits the continuous exposure to infectious diseases, there is evidence suggesting that opportunities for disease transmission have occurred (Lamberski et al., 2009:243-245). However, the results are limited to a small number of BFC tested, as well as being from only a few locations, and as such, cannot be inferred to all BFC across their range. More information is needed about the genetic background and immune competence of BFC, both spatially and temporally. What is known is that contagious multihost viruses such as canine distemper is more easily spread and result in animals either dying or obtaining a lifelong immunity (Craft, Hawthorne, Packer & Dobson, 2008:1262).

The discovery of AA-amyloidosis in wild BFC suggests a predilection for the disease. Originally considered a stress-induced disease in captive cats, it is now suspected that wild BFC experiencing any forms of intrinsic or extrinsic stress may also develop the disease. It is a priority of the BFCWG to repeatedly test and monitor subject cats in an attempt to determine onset and possible causes for the

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development of the disease. It is also hoped to establish whether AA-amyloidosis disease-free populations of BFC exist in the wild, which may be used to bolster or replace captive populations.

The long-term effects of climate change and global warming cannot be overlooked. Changes in weather patterns may lead to changes in range; in the timing of breeding events; increases in severe weather such as flooding and droughts; altered disease patterns, or increased risks of the spread of pathogens from parasites. Of course, dry and hotter climates may result in increasing termite ranges. This may be beneficial to BFC particularly in Botswana and elsewhere in the northern areas of its range. However, expanding termite ranges may result in increased pest damage, particularly during drought periods, which will also be more frequent. This will have huge economic implications for the subsistence farmers in those regions and will stimulate a need for termite control policies to be implemented (Nkunika et al., 2010).

Overall, climate change is more often predicted to have rapid and severe negative influences on all apex predator abundance rather than benefits. The rate of change often exceeds the ability of animals to adapt or disperse to more climatically favourable regions where prey densities are sufficient to support viable populations. Black-footed cats have already adapted to changing habitats over the past two million years, but the ability to adapt further may depend where they are in their evolutionary lifespan, as well as their genetic and behavioural resilience that remains undetermined at this point.

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5.5.4 Ranking of the threats

In line with most conservation actions, and during the recent review for the conservation status of BFC for The Red List of Mammals of South Africa,

Swaziland and Lesotho (Wilson, Sliwa & Drouilly, in prep.), threats are ranked and described to correlate with threats facing other species and are linked to the scale of those threats throughout the ranges of the species. Severity refers to whether the threat is reversible and can be managed either at a metapopulation level, or aimed at extrinsic factors that involve multiple stakeholders. The previous risks listed above are summarised and ranked in Table 5.3 based on the findings by

Wilson, Sliwa & Drouilly (in prep.).

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TABLE 5.3: A summary of the conservation threats to black-footed cats (Felis nigripes) ranked in order of decreasing severity

Data type: Time Scale: Trend:

Period: observed,

Threat description local, estimated, stable, Severity past, Rank regional, projected, increasing, present, national inferred, decreasing future suspected

Highly fragmented and patchy Observed, 1 distribution with Regional All Increasing Unmanageable Suspected resulting genetic and dispersal implications

Human induced habitat degradation / Stable, loss resulting in loss Observed, 2 Regional All possibly Unmanageable of prey base and inferred increasing other resources (e.g. springhares)

Absence from Stable, National & Observed, 3 formally protected All possibly Manageable Regional inferred areas decreasing

Moderate to high Stable, Observed, 4 black-backed jackal Local All possibly Manageable suspected and caracal numbers increasing

Persecution through accidental poisoning through pesticides or 5 Local Suspected All Stable Unmanageable poison intended for other problem species

Victimisation by feral, 6 hunting and guarding Local Suspected All Increasing Manageable dogs

National & 7 Road mortalities Observed All Increasing Unmanageable Regional

Observed, 8 Natural calamities Local All Stable Unmanageable suspected

Stable, Diseases (inherited National & Observed, 9 All possibly Unmanageable and transmitted) Regional suspected increasing

Trophy industry Observed, 10 National All Increasing Manageable exploitation suspected

11 Climate change Regional Suspected All Increasing Unmanageable

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5.6 SUMMARY

The historical paucity of data for the BFC has led to inconsistencies in previous determinations of the geographical range, prediction of population size and the potential threats the species is facing. This study has proposed a repeatable method for estimating the population size of BFC when conducting conservation assessments. Using this method, the current estimations of mature population size suggest 8 333 individuals in South Africa and 9 707 in the entire regional population, with no subpopulation consisting of more than 1 000 individuals.

However, the results may be severely confounded by reporting biases and the lack of regional spatial and temporal data in the regions outside of South Africa.

The species is currently only afforded formal protection status in Botswana and

South Africa. In particular, the conservation status of BFC in Namibia needs to be determined and adequate conservation measures implemented. After reviewing the known conservation statuses throughout the region, as well as the previously identified threats, 13 potential threats were identified. The majority of the threats facing BFC have been observed during this study and throughout the research underway by the BFCWG. However, some of the threats need to be investigated further and quantified.

The findings of this study suggest that BFC were not threatened by possible hybridisation with other cat species. The species appears marginally more widespread and utilises more habitat types than previously documented. There is the suggestion that the species may benefit from judicious predator control

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measures, particularly in the central, sheep-farming areas of South Africa. Both the BFC and this research benefited from the increased public awareness of the species.

However, the threats faced by BFC appear significantly more increased and varied than previously documented. Habitat fragmentation, disturbances and degradation together have the potential to lead to other threats such as disruption of dispersal, population genetics, loss of prey base and the reduction in available safe refuges. They also lead to increased levels of stress, exposure to contagious diseases, and vulnerability to intraguild predation. The true impact of road mortalities and the trophy hunting industry are not known, but evidence suggests that both aspects are taking a toll to some degree on the species. The long-term effects of global warming need to be considered as this will alter significant habitats and prey base composition.

After reviewing all the threats, 11 were determined to have either intrinsic or anthropogenic risks to BFC. These were summarised and ranked according to the scale, data type, time period, trend and severity of the threat.

Unfortunately, the majority of the threats appear to be increasing and unmanageable, and the disturbing low reporting rate from formally protected areas may indicate the species to be absent in these areas which is a cause for concern.

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CHAPTER 6

CONCLUSION AND RECOMMENDATIONS

6.1 INTRODUCTION

Carnivores, as apex predators, are biodiversity indicators of ecosystems.

Information about their historical and present distributions together with estimations of population size may indicate what changes have occurred to the ecosystems in which they live. In turn, this information also assists with the conservation assessment for a particular species by highlighting the threats facing the species. This then allows for the formation of appropriate management recommendations and research priorities going forward.

Until recently, the historical paucity of data on the black-footed cat (BFC) (Felis nigripes) has led to inconsistencies and perpetuated inaccuracies in current literature. This, to date, has affected the accuracy of conservation measures and management recommendations for the species at both a national and regional level.

6.2 SYNTHESIS OF THE FINDINGS

6.2.1 Geographical distribution

The reliability and validity of data contributed by citizen scientists is often criticised.

However, careful vetting of the data and personalised interview processes can

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greatly eliminate these problems, and there are substantial benefits to having had members of the public involved. The area covered and the number of records obtained is increased at very little cost to the researcher. It also served to demonstrate to the citizen scientists that their participation serves national and international biodiversity goals. This study also created opportunities for them to articulate these understandings within their communities, which in turn, helped create a broader awareness of BFC in their areas. This was considered particularly important once it was discovered that the species is offered only marginal protection in formally protected areas and the possible future of the species lies in the hands of private landowners.

Despite examining and mapping 790 historical and current locality records, the true range of BFC is still impossible to map with complete accuracy due to the lack of spatial and temporal data. By mapping the marginal occurrence records, a new geographical distribution of 2 214 267 km2 for the species is proposed. This map is suitable for use in standard field guides. Although this map is nearly twice that of the first range estimations of Stuart in1982, the species still has a highly restricted range. The new range extensions were primarily north- and eastwards. The species remains absent from Lesotho and Swaziland, but is now believed to be at least marginally present in southern Angola and western Zimbabwe.

The weak sampling of localities and the lack of spatial and temporal data from

Namibia and Botswana, and possibly even from Angola and Zimbabwe, may result in significant errors of omission and commission. This could undermine accurate population size estimations for each country and the region as a whole.

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The new EOO map appears to have narrowed the errors of omission that were evident in historical maps and increased the overall range. However, this range extension cannot be inferred as a true increase in the geographical distribution, but rather as an artefact of historical under-reporting and lack of research effort.

6.2.2 Conservation status

The current estimations of BFC population size suggest there to be 8 333 mature individuals in South Africa and 9 707 regionally. Previous inaccuracies in population size estimations had the species incorrectly listed only as Least

Concern in South Africa. Based on the results of this study, this listing has recently been revised to Vulnerable (Wilson, Sliwa & Drouilly, in prep.). This is also the current global conservation status, and both assessments are considered appropriate based on the evidence herein.

Both natural and anthropogenic factors are important in determining a species’ risk of extinction and priority needs for conservation measures. A variety of both intrinsic traits and extrinsic anthropogenic factors may be placing BFC at a higher risk of expedited population declines. The findings herein suggest that most risks have gone undetected during past conservation assessments. This is probably due to a lack of consistent research and reporting efforts.

Habitat-changing factors were considered the most significant threats as these leads to other potential threats developing or exacerbate those already present.

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Unfortunately, the BFC also faces a number of intrinsic health and genetic issues that may affect its overall ability to adapt to a changing environment.

Monitoring of the species remains challenging due to its rarity and elusive nature.

They are consistently recorded in literature as rare and elusive, and this situation is unlikely to change, even with public awareness. Investigations into the degree of dependence on sympatric species may warrant further investigation. An easily seen and correctly identified species such as a springhare may prove to be a good surrogate species for predicting BFC distribution in areas where reporting rates would otherwise be low due to a lack of familiarity with the cats.

After ranking the severity of the 11 identified threats in terms of scale, type, time period and trend, the majority of the threats appear to be increasing and unmanageable.

6.3 CONSERVATION IMPLICATIONS

Studies are urgently required especially on the small, less charismatic and lesser- known carnivores (Keith, 2005:40). To date there is a severe and documented conservation bias towards larger more charismatic taxa that sees them being awarded the bulk of funding and attention (Polishchuk, 2002:1123).

Raising the conservation status of the species will hopefully increase research interest and access to funding. Increased research effort on this species will contribute towards a better understanding of the needs of the species and the

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threats it is facing. All the findings of this study imply that the species is facing more threats than previously realised, quite possibly a declining population size in an increasingly fragmented habitat. The causes are varied but anthropogenic factors are probably having the greatest impact but which, in turn, can exacerbate certain key intrinsic features of the species.

Human-initiated habitat changes that have resulted in a highly fragmented and patchy distribution may be creating local genetic bottlenecks, increasing susceptibility to diseases, and reducing reproductive fitness. Reports of cats in formally protected areas are relatively rare and these areas need to prioritise population censuses for the species. Although these areas are normally too small to support a full subpopulation of BFC, it would be reassuring to have some protected habitats available for the species. In addition, private landowners in suitable habitat could be targeted to champion the species by limiting habit changes and employing better farming practices management strategies that address the increasing problem predator numbers. Such areas have the potential to preserve (or reopen) the travel corridors that cats use to move from one area to another.

The continued well-being of BFC may also be, in part, inextricably linked or dependent on other non-prey species. For instance, springhares are not a protected species and are currently, globally and nationally, listed as Least

Concern. Although widespread and abundant in some areas, they are in a rapid decline over much of their geographic range due to habitat degradation, habitat loss, and hunting (Butynski, 2013:623). The springhare is likely to be a keystone

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species in some ecosystems and as such, probably needs to have its conservation status reassessed, not only for the long-term survival of the species but also for other species with close associations such as BFC. Other species that have an important impact of the continued survival of BFC are intraguild species such as black-backed jackals, caracals and herding dogs that have had a proven impact on the species and, if left unchecked, may account for as much as 50% of the mortalities in a local cat population.

With limited resources available for conservation, it is generally accepted that if a species is on the Red List it should be a conservation priority. The identification of a species that demands special conservation measures or which needs to be a regional priority provides invaluable information for the execution of conservation plans.

South Africa’s Red Data Lists are some of the most complete amongst the African countries. This study has served to illuminate the potential shortfall of suitable data from neighbouring African countries. This is particularly crucial in the case of the

BFC as regional information usually feeds into global assessments. The BFC is currently only afforded formal protection status in South Africa and Botswana, although the species has a known presence in Namibia. Other countries urgently need to establish formalised conservation assessment programmes too.

In addition, these assessments need to take place regularly. Previous assessments, in South Africa for instance, have had long lapses between reviews

(Smithers, 1986; Friedman & Daly, 2004). The current assessment underway is

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taking place after just a 10-year interval resulting in a significant uplifting of the conservation status of BFC.

6.4 MANAGEMENT RECOMMENDATIONS

Noss (1992:11), the internationally renowned conservation biologist, once said, “In conservation, science tells us what to do; but our values tell us why we should do it”. Whilst the species may not be a keystone species in its environment, it still plays an important ecosystem role as a dominant predator of small mammals and birds in the areas they inhabit.

In line with most conservation actions and the recent review for the conservation status of the species for The Red List of Mammals of South Africa, Swaziland and

Lesotho (Wilson, Sliwa & Drouilly, in prep.), a number of conservation interventions are suggested for the entire region and ranked in order of priority.

These actions are linked to the scale of the intervention throughout the range of the species. The basis of the ranking is indicated on how the threat was determined, as well as whether the action suggested is already in place or needs to be implemented in the future. Lacking further information at this point, success rates are mostly unknown. A summary of some of the possible conservation interventions are given in Table 5.3 based on the national findings by Wilson,

Sliwa and Drouilly (in prep.). With future research, these interventions can be reviewed and adapted as additional information becomes available.

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TABLE 6.1: A summary of proposed conservation interventions for the black- footed cat (Felis nigripes) in order of priority Data type: Scale: observed, In place, Success rate: local, estimated, planned or Intervention description % target national, projected, future Rank achievement regional inferred, potential? suspected Judicious management and In place in reduction of unnaturally high Observed, some areas 1 jackal and caracal numbers Local Unknown suspected but needs to via human control be extended Education and awareness of landowners to reduce Future 2 Local Suspected Unknown accidental killings and habitat potential degradation Establishment of suitably Local & Future 3 large conservancy areas Suspected Unknown Regional potential (<100 000 hectares) Maintenance of viable local resources (e.g. springhare Local & Future 4 Suspected Unknown populations for burrow Regional potential refuges, etc.) Public awareness to raise National & Future 5 Suspected Unknown profile of this cryptic species Regional potential Raising the conservation status means stiffer penalties In place, can be enforced on National & 6 Suspected future Unknown transgressors deliberately Regional potential targeting or accidentally killing cats Creation of a management plan for the species as an National Future 7 Suspected Unknown interim/pre-emptive Regional potential conservation measure In place, Ex situ management National 8 Observed future Limited interventions Regional potential

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6.5 FUTURE RESEARCH PRIORITIES

The BFC has been extensively studied for more than 20 years near the Kimberley area, along the Free State-Northern Cape provincial border in central South Africa.

This has provided information about the diet (Sliwa, 2006:195-204); home range size and social organisation (Sliwa, 2004:96-107); reproductive biology (Herrick et al., 2010:552-562); diseases (Lamberski et al., 2009:243-245); ecological relationships between the species and other sympatric carnivores (Kamler et al.,

2015:122-127), and various other findings (Sliwa et al., 2007; Sliwa et al., 2008;

Sliwa et al., 2009a; Sliwa et al., 2009b; Sliwa et al., 2010; Sliwa et al., 2011; Sliwa et al., 2013; Sliwa et al., 2014; Sliwa et al., 2015).

More recently, the BFCWG extended the focus of its field research to include farms south of De Aar, Northern Cape, South Africa, which have different vegetation types with differing farming practices in place. However, there is still a paucity of information for BFC elsewhere within its range and the following are considered some of the research priorities by Wilson, Sliwa and Drouilly (in prep.):

 Fine scale distributional studies across the entire distributional range.

 Accurate subpopulation estimates and long-term monitoring of subpopulation

trends are needed throughout the species geographical range.

 The impacts and extent of persecution, both direct and indirect on

subpopulations needs to be established.

 The effects and impacts of apex carnivores on BFC populations need to be

determined.

 The effects of long-term geographical isolation on subpopulations.

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 Investigation into the validity of the two possible subspecies and the resulting

conservation implications.

 Studies on changes in density across a spectrum of habitat quality.

 Surveys to reveal the presence/absence of the species in formally protected

areas.

 Studies into the dispersal abilities and survival of subadult BFC in differing

habitats types.

 Fine-scale determination on the habitat characteristics and prey populations

characteristics required by female BFC to successfully raise kittens.

 Investigation into the causes of kitten mortality and survival rates.

 Determination of the extent of direct persecution by farmers and the efficacy of

education and awareness programmes targeted at landowners.

 Investigation into the causes and extent of AA-amyloidosis in wild populations.

 The degree of dependence on sympatric species e.g. hollow termitarium for

kittens and springhares for burrow refuges, and the effects on the long-term

survival of BFC following the removal of host populations.

 The range and impact of transmittable diseases from sympatric carnivores on

BFC.

 The refinement of artificial insemination techniques using cryopreserved sperm

from BFC.

 The refinement of the procedures and techniques of in vitro fertilisation in BFC.

 Determination of the numbers of BFC being illegally removed for the trophy

industry.

 Monitoring of the success of re-introduced individuals into new areas

with/without other individuals present.

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 Monitoring of population trends.

 Establishing the effects of electric fences on local movements and dispersal

patterns of BFC.

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REFERENCES

AKERLOF, K. & KENNEDY, C. 2013. Nudging toward a healthy natural environment – how behavioural change research can inform conservation.

Fairfax: George Mason University.

ANGOLA. Department of Environment, 2009. Framework report on Angola’s

Biodiversity. Fourth National Angola report to the Convention on Biological

Diversity. Luanda.

AVENANT, N. & CAVALLIN, P. 2007. Correlating rodent community structure with ecological integrity, Tussen-die-Riviere Nature Reserve, Free State province,

South Africa. Integrative Zoology 2:212-219.

BAGNIEWSKA, J.M. & KAMLER, J.F. 2013. Do black-backed jackals affect numbers of smaller carnivores and prey? African Journal of Ecology 52:564-567.

BAMFORD, M.K., NEUMANN, F.H., PEREIRA, L.M., SCOTT, L., DIRKS,

P.H.G.M. & BERGER, L.R. 2010. Botanical remains from a coprolite from the

Pleistocene hominin site of Malapa, Sterkfontein Valley, South Africa.

Palaeontologica Africana 45:23-28.

BOONEY, R, COOPER, C.B., DICKINSON, J., KELLING, S., PHILLIPS, T.,

ROSENBERG, K.V. & SHIRK, J. 2009. Citizen science: a developing tool for expanding science knowledge and scientific literacy. BioScience 39 (11):977-984.

143

BOSHOFF, A.F. & KERLEY, G.H. 2010. Historical mammal distribution data: how reliable are written records? South African Journal of Science 106,1/2:26-33.

BOTSWANA. Ministry of Environment, Wildlife and Tourism, 2009. Fourth

National Botswana report to the Convention on Biological Diversity.

Board on Science and Technology for International Development National

Research Council (BOSTID). 1991. Microlivestock: Little-known Small Animals with a Promising Economic Future. National Academy of Sciences. Washington

DC: National Academy Press.

BRODIE, J.F. 2009. Is research effort allocated efficiently for conservation?

Felidae as a global case study. Biodiversity and Conservation 18:2927-2939.

BROMAN, D.J.A., LITVAITIS, J.A., ELLINGWOOD, M., TATE, P. & REED, G.C.

2014. Modeling bobcat Lynx rufus habitat associations using telemetry locations and citizen-scientist observations: are the results comparable? Wildlife Biology

20:229-237.

BRONNER, G.N., HOFFMAN, M., TAYLOR. P.J., CHIMIMBA, C.T., BEST, P.B.,

MATTHEE, C.A. & ROBINSON, T.J. 2003. A revised systematic checklist of the extant mammals of the southern African subregion. Durban Museum Novitates 28:

56-106.

144

BROOKE, Z.M., BIELBY, J., NAMBIAR, K. CARBONE, C. 2014. Correlates of research effort in carnivores: body size, range size and diet matter. PLoS ONE

9(4): e93195.

BURCHELL, W.J. 1824. Travels to the Interior of Southern Africa, vol. 2.

London: Longman, Hurst, Rees, Orme, Brown and Green.

BUTYNSKI, T.M. 1973. Life history and economic value of the springhare

(Pedetes capensis Forster) in Botswana. Botswana Notes and Records 5:200-213.

BUTYNSKI, T.M. 2013. Pedetes capensis Southern African Springhare. In:

Happold, D.C.D. (ed.). Mammals of Africa, vol. 3: rodents, hares and rabbits.

London: Bloomsbury Publishing.

COFFIN, A.W. 2007. From roadkill to road ecology: a review of the ecological effects of roads. Journal of Transport Geography 15:396-406.

COLLINSON, W. 2014. Telephonic discussion. Wildlife and Roads Project,

Endangered Wildlife Trust. Tel. +27 11 172 3600. Email: [email protected]

CRAFT, M.E., HAWTHORNE, P.L., PACKER, C. & DOBSON, A.P. 2008.

Dynamics of a multihost pathogen in a carnivore community. Journal of Animal

Ecology 77:1257-1264.

145

DAVIS, H.A. S.a. Precious samples provide insight into black-footed cat genetics

[Online]. Available from: http://www.sandiegozooglobal.org/what_we_do_preserving_wildlife/mammals/prec ious_samples_provide_insight_into_black-footed_cat_genetics/ [Accessed:

15/4/2015].

DAVIS, A.R. & GRAY, D. 2010. The distribution of Scottish wildcats (Felis silvestris) in Scotland (2006-2008). Scottish Natural Heritage Commissioned

Report No. 360 [Online]. Available from: www.snh.org.uk/pdfs/publications/commissioned_reports/360.pdf [Accessed:

15/4/2015].

DICKSON, J.L., ZUCKERBERG, B. & BONTER, D.N. 2010. Citizen science as an ecological research tool: challenges and benefits. Annual Review of Ecology,

Evolution, and Systematics 41:149-172.

DO LINH SAN, E. & SOMERS, M.J. 2013. Editorial – African small carnivores: the

‘forgotten Eden’. Small Carnivore Conservation 48:1-2.

DO LINH SAN, E., FERGUSON, A.W., BELANT, L., SCHIPPER, J., HOFFMANN,

M. GAUBERT, P. ANGELICI, M. & SOMERS, M.J. 2013. Conservation status, distribution and species richness of small carnivores in Africa. Small Carnivore

Conservation 48:4-18.

146

DURRHEIM, K. 2006. Quantitative measurement. In: Terre Blanche, M. &

Durrheim, K. (eds.). Research in practice – applied methods for the social sciences. Cape Town: University of Cape Town Press.

ESRI. 2011. ArcGIS Desktop: Release 10. Redlands, CA: Environmental

Systems Research Institute. Available from: http://www.esri.com/software/arcgis/new

FORDHAM, D.A., AKCAKAYA, H.R., BROOK, B.W., RODRIGUEZ, A., ALVES,

P.C., CIVANTOS, E., TRIVINO, M., WATTS, M.J. & ARAUJO, A.B. 2013.

Adapted conservation measures are required to save the Iberian lynx in a changing climate. Nature Climate Changes 3:899-903.

FERRAR, P. 1982. The termites of the Savanna Ecosystem Project, study area,

Nylsvley. South African National Scientific Programmes Report 60, September

1982:1-41. Pretoria: CSIR.

FRIEDMAN, Y. & DALY, D. (eds.). 2004. Red Data Book of the Mammals of

South Africa: a conservation assessment. Johannesburg: Conservation Breeding

Specialist Group (SSC/IUCN) & The Endangered Wildlife Trust.

GASTON, K.J. & FULLER, R.A. 2009. The sizes of species’ geographic ranges.

Journal of Applied Ecology 46:1-9.

147

GELDERBLOM, C.M., BRONNER, G.N., LOMBARD, A.T. & TAYLOR, P.J. 1995.

Patterns of distribution and current protection status of the Carnivora, Chiroptera and Insectivora in South Africa. South African Journal of Zoology 30(3):103-114.

GUISAN, A. & ZIMMERMAN, N.E. 2000. Predictive habitat distribution models in ecology. Ecological Modelling 135:147-186.

HAUB, C. & KANEDA, T. 2013. 2013 World population data sheet. Washington

DC: Population Reference Bureau.

HERRICK, J.R., CAMPBELL, M., LEVENS, G., MOORE, T., BENSON, K.,

D’AGOSTINO, J., WEST, G., OKESON, D.M., COKE, R., PORTACIOS.C.,

LEISKE, K., KREIDER, C., POLUMBO, P.J. & SWANSON, W.F. 2010. In vitro fertilization and sperm cryopreservation in the black-footed cat (Felis nigripes) and sand cat (Felis margarita). Biology of Reproduction 82:552-562.

HORACK, I.G., HEYNE, H. & DONKIN, E.F. 2010. Parasites of domestic and wild animals in South Africa. XLVIII. Ticks (Acari: Ixodidae) infesting domestic cats and wild felids in southern Africa. Onderstepoort Journal of Veterinary Research 77(1):

Art. #3.

HOSKIN, C., HIGGIE, M., McDONALD, K.R. & MORITZ, C. 2005. Reinforcement drives rapid allopatric speciation. Nature 437 (7063):1353-1356.

148

IUCN. 2001. IUCN Red List Categories and Criteria: Version 3.1. IUCN Species

Survival Commission, IUCN, Gland, Switzerland and Cambridge, UK. Oxford:

Information Press.

IUCN / SSC CAT SPECIALIST GROUP. 2015. Cat Species Information

[ONLINE]. Available from: http://www.catsg.org/index.php?id=4 [Accessed:

30/4/2015].

JETZ, W., SERERCIOGLU, C.H. & WATSON, E.M. 2008. Ecological correlates and conservation implications of overestimating species geographic ranges.

Conservation Biology 22:110-119.

JOUBERT, E., MORSBACH, D. & WALLIS, V. 1982. The 1982 distribution patterns and status of some mammals on farms in South West Africa.

Unpublished Report. Departmental Report N 18/2/21, Directorate of Nature

Conservation & Recreational Resorts, Windhoek.

KAMLER, J.F., STENKEWITZ, U., SLIWA, A., WILSON, B., LAMBERSKI, N.,

HERRICK, J.R. & MACDONALD, D.W. 2015. Ecological relationships of black- footed cats (Felis nigripes) and sympatric canids in South Africa. Mammalian

Biology 80:122-127.

KEITH, M. 2005. Conservation assessment of South African mammals. PhD thesis, Pretoria, University of Pretoria.

149

KLEIN, R.G., CRUZ-URIBE & BEAUMONT, P.B. 1991. Environmental, ecological and paleoanthropological implications of the Late Pleistocene mammalian fauna from Equus Cave, Northern Cape Province, South Africa. Quaternary Research

36:94-119.

KUHN, B.F., WERDELIN, L., HARTSTONE-ROSE, A., LACRUZ, R.S. &

BERGER, L.R. 2011. Carnivore remains for the Malapa Hominin site, South

Africa. PLoS ONE 6(11): e26940.

KUHN, B.F. 2013. Interview. Institute for Human Evolution, University of the

Witwatersrand, Johannesburg, South Africa.

LAMBERSKI, N. S.a. Stepping up to save black-footed cats [ONLINE]. Available from: www.sandiegozooglobal.org/what_we_do_preserving_wildlife/mammals/stepping_ up_to_save_black-footed_cats/. [Accessed: 5/4/2015].

LAMBERSKI, N., SLIWA, A., WILSON, B., HERRICK, J., LAWRENZ, A. &

DUBOVI, E. 2009. Conservation of black-footed cats and prevalence of infectious diseases in sympatric carnivores in the Northern Cape province, South Africa. In:

WIBBELT, G., KRETZSCHMAR, P., HOFER, H. & SEET, S. (eds.). Proceedings of the International Conference on diseases of zoo and wild animals organised by

Leibniz Institute for Zoo and Wildlife Research, Berlin, Germany. Liebefeld-Berne:

European Association of Zoo and Wildlife Veterinarians:243-245.

150

LEIRS, H., STENSETH, N.C., NICHOLS, J.D., HINES, J.E., VERHAGEN, R. &

VERHEYEN, W. 1997. Stocastic seasonality and nonlinear density-dependent factors regulate population size in an African rodent. Nature 389:176-180.

LE ROUX, J.J., FOXCROFT, L.C., HERBST, M. & MACFADYEN, S. 2015.

Genetic analysis shows low levels of hybridization between African wildcats (Felis silvestris lybica) and domestic cats (F. s. catus) in South Africa. Ecology and

Evolution 5(2):288-299.

LEYHAUSEN, P. 1979. Cat Behaviour: The predatory and social behaviour of domestic and wild cats. Translated by B.A. Tonkin: Katzen, eine Verhaltenskunde.

1975. New York: Garland STPM Press.

LINDSEY, P., DU TOIT, J.T. & MILLS, M.G.L. 2004. The distribution and population status of African wild dogs (Lycaon pictus) outside protected areas in

South Africa. S.A. Journal of Wildlife Research 34(2):143-151.

LIVINGSTONE, D. 1857. Missionary travels and researches in South Africa, vol. 1.

London: Murray.

LOPEZ, J.V., CULVER, M., STEPHENS, C., JOHNSON, W.E. & O’BRIEN, S.J.

1997. Rates of nuclear and cytoplasmic mitrochondrial DNA sequence divergence in mammals. Molecular Biology and Evolution 14(3):277-286.

151

LOVERIDGE, A.J., WANG, S.W., FRANK, L.G. & SEIDENSTICKER, J. 2010.

People and wild felids: conservation of cats and management of conflicts. In:

Macdonald, D.W. & Loveridge, A.J. (eds.). Biology and Conservation of Wild

Felids. Oxford: University Press:161-195.

LYNCH, C.D. 1975. The distribution of mammals in the Orange Free State, South

Africa. Navorsing van die Nasionale Museum, Bloemfontein 3:109-139.

MACDONALD, D.W., LOVERIDGE, A.J. & NOWELL, K. 2010. Dramatis personae: and introduction to the wild felids. In: Macdonald, D.W. & Loveridge,

A.J. (eds.). Biology and Conservation of Wild Felids. Oxford: University Press:3-

58.

MATTHEE, S., VAN DER MESCHT, L., WILSON, B. & LAMBERSKI, N. 2011.

Flea diversity on small carnivores in the Northern Cape province, South Africa.

African Zoology 46(1):27-31.

MCBRIDE, D.M. 2013. The process of research in psychology. London: Sage.

MCKELVEY, K.S., AUBRY, K.B. & SCHWARTZ, M.K. 2008. Using anecdotal occurrence data for rare or elusive species: the illusion of reality and a call for evidentiary standards. BioScience 58(6):549-555.

152

MCMANUS, J.S., DALTON, D.L., KOTZE, A., SMUTS, B., DICKMAN, A.,

MARSHAL, J.P. & KEITH, M. 2015. Gene flow and population structure of a solitary top carnivore in a human-dominated landscape. Ecology and Evolution

5(2):335-344.

MEESTER, J.A.J., RAUTENBACH, I.L., DIPPENAAR, N.J. & BAKER, C.M. 1986.

Classification of Southern African Mammals. Transvaal Museum Monograph 5:1-

359.

MILLS, L.S. 2007. Conservation of wildlife populations: demography, genetics and management. Malden: Blackwell Publishing.

MINISTRY of Environment see ANGOLA.

MINISTRY of Environment, Wildlife and Tourism see BOTSWANA.

MINISTRY for the Coordination of Environmental Affairs see MOZAMBIQUE.

MINISTRY of Environment and Tourism see NAMIBIA.

MINISTRY of Tourism, Environment and Natural Resources see ZAMBIA.

MINISTRY of Environment & Natural Resources Management see ZIMBABWE.

153

MOLTENO, A.J., SLIWA, A. & RICHARDSON, P.R.K. 1998. The role of scent marking in a free-ranging, female black-footed cat (Felis nigripes). Journal of

Zoology 245:35-41.

MOZAMBIQUE. Ministry for the Coordination of Environmental Affairs, 2014.

Fifth National Report on the Implementation of Convention on Biological Diversity in Mozambique. Maputo.

NAMIBIA. Ministry of Environment and Tourism, March 2014. Fifth National

Report to the Convention on Biological Diversity (2010-2014).

NATTRASS, N. & CONRADIE, B. 2013. Jackal narratives and predator control in the Karoo, South Africa. CSSR Working Paper No. 324, June 2013. Cape Town:

Centre for Social Science Research, UCT.

NKUNIKA, P.O.Y., SHIDAY, B.M.A., SILESHI, G.W., FRENCH, J.R.J., NYEKO, P.

& JAIN, S. 2010. Termite taxonomy and distribution with particular reference to climate change in Africa. (Paper read at The International Research Group on

Wood Protection, 09 May 2010, Biarritz, France). Unpublished. Available from: www.outputs.worldagroforestry.org/record/5570/files/PP11012.pdf

NOSS, R.F. 1992. The Wildlands Project: land conservation strategy. Wild Earth

(special issue):10-25.

154

NOWELL, K. & JACKSON, P. (eds.). 1996. Wild Cats: Status Survey and

Conservation Action Plan. Switzerland: IUCN.

NUIJTEN, E. 2011. Combining research styles of the natural and social sciences in agricultural research. Wageningen Journal of Life Sciences 57:197-205.

OLBRICHT, G. & SLIWA, A. 1997. In situ and ex situ observations and management of black-footed cats (Felis nigripes). In: Olney, P.J.S. and Fisken,

F.A. International Zoo Yearbook 35: Felids. London: The Zoological Society of

London:81-89.

PALMA, L., BEJA, P. & RODRIGUES, M. 1999. The use of sighting data to analyse Iberian lynx habitat and distribution. Journal of Applied Ecology 36:812-

824.

PEARSON, R.G., RAXWORTHY, C.J., NAKAMURA, M. & PETERSON, T. 2007.

Predicting species distributions from small numbers of occurrence records: a test case using cryptic geckos in Madagascar. Journal of Biogeography 34(1):102-117.

PETERS, G. & BAUM, L., PETERS, M.K. & TONKIN-LEYHAUSEN, B. 2007.

Spectral characteristics of intense mew calls in cat species of the genus Felis

(Mammalia: Carnivora: Felidae). Journal of Ethology 27:221-337.

155

PLUG, I. & BADENHORST, S. 2001. The distribution of macromammals in southern Africa over the past 30 000 years as reflected in animal remains from archaeological sites. Transvaal Museum Monograph 12:1-.234

POCOCK, R.I. 1907. Notes upon some African species of the genus Felis, based upon specimens recently exhibited in the Society Gardens. Proceedings of The

Zoological Society of London 1907:656-677.

POLISHCHUK, L.V. 2002. Conservation priorities for Russian mammals. Science

297:1123.

POTGIETER, G.C., MARKER, L.L., AVENANT, N.L. & KERLEY, G.I.H. 2013. Why

Namibian farmers are satisfied with the performance of their livestock guarding dogs. Human Dimensions of Wildlife: An International Journal 18(6):403-415.

PURVIS, A., GITTLEMAN, J.L., COWLISHAW, G. & MACE, G.M. 2000.

Predicting extinction risk in declining species. Proceedings of the Royal Society of

London 267:1947-1952.

QUANTUM GIS DEVELOPMENT TEAM (QGIS). 2015. QGIS Geographic

Information System. Open Source Geospatial Foundation Project. Available from: http://qgis.osgeo.org

156

RAMESH, T., KALLE, R., SANKAR, K. & QURESHI, Q. Spatio-temporal partitioning among large carnivores in relation to major prey species in Western

Ghats. Journal of Zoology 287(4):269-275.

RAUTENBACH, I.L. 1982. Mammals of the Transvaal. Ecoplan Monograph

1:111-211.

RONDININI, C. & BOITANI, L. 2012. Mind the map: trips and pitfalls in making and reading maps of carnivore distribution. In: Boitani, L. and Powell, R.A. (eds.).

Carnivore Ecology and Conservation. New York: Oxford University Press.

ROWE-ROWE, D.T. 1992. The carnivores of Natal. Pietermaritzburg: Natal Parks

Board.

SA see SOUTH AFRICA.

SCHIPPER, J., HOFFMANN, M., DUCKWORTH, J.W. & CONROY, J. 2008. The

2008 IUCN red listings of the world’s small carnivores. Small Carnivore

Conservation 39:29-34.

SHEA, J. 2014. Small wild cats face big threats, but receive little conservation funds. Earth Island Journal. [Online]. Available from: http://www.earthisland.org/journal/index.php/elist/eListRead/small_wild_cats_face_ big_threats_but_receive_little_conservation_funds/

157

SHORTRIDGE, G.C. 1931. Felis (Microfelis) nigripes thomasi subsp. nov.

Records of the Albany Museum 4:1.

SHORTRIDGE, G.C. 1934. The Mammals of South West Africa: A biological account of the forms occurring in that region, vol. 1. London: Heinemann.

SKINNER, J.D & CHIMIMBA, C. (eds.). 2005. The mammals of the southern

African subregion. 3rd edition. Cambridge: Cambridge University Press.

SKINNER, J.D. & SMITHERS, R.H.N. 1990. The Mammals of the Southern

African Subregion. 2nd ed. Pretoria: University of Pretoria Press.

SLIWA, A. 1994. Diet and feeding behaviour of the black-footed cats (Felis nigripes Burchell, 1824) in the Kimberley Region, South Africa. Der Zoologische

Garten (n.f.) 64:83-96.

SLIWA, A. 2004. Home range size and social organisation of black-footed cats

(Felis nigripes). Mammalian Biology 69 (2):96-107.

SLIWA, A. 2006. Seasonal and sex-specific prey composition of black-footed cats Felis nigripes. Acta Theriologica 51 (2):195-204.

SLIWA, A. 2008. Felis nigripes. The IUCN Red List of Threatened Species.

Version 2014.2 [Online]. Available from: www.iucnredlist.org [Accessed:

15/09/2014].

158

SLIWA. A. 2013. Felis nigripes Black-footed Cat. In: Kingdon, J. & Hoffman, M.

(eds.). Mammals of Africa vol. 5: carnivores, pangolins, equids and rhinoceroses.

London: Bloomsbury Publishing:203-206.

SLIWA, A. 2014. Telephonic conversation. Black-footed Cat Working Group

Project Leader; Curator, Cologne Zoo, Germany. Tel. +49-221-7785-195.

Fax: +49-221-7785-111. Email: [email protected]

SLIWA, A. 2015. Email correspondence. Black-footed Cat Working Group Project

Leader; Curator, Cologne Zoo, Germany. Tel. +49-221-7785-195. Fax: +49-221-

7785-111. Email: [email protected]

SLIWA, A., HERBST, M. & MILLS, M.G. 2010. Black-footed cats (Felis nigripes) and African wildcats (Felis silvestris): a comparison of two small felids from South

African arid lands. In: Macdonald, D.W. & Loveridge, A.J. (eds.). Biology and

Conservation of Wild Felids. Oxford: University Press:535-558.

SLIWA, A., WILSON, B., KUSTERS, M. & TORDIFFE, A. in prep. Felis nigripes.

The IUCN Red List of Threatened Species. Version 2016-?. [Online]. Available from: www.iucnredlist.org.

SLIWA, A., WILSON, B., LAMBERSKI, N. AND HERRICK, J. 2007. Report on catching and surveying black-footed cats (Felis nigripes) on Benfontein Game

Farm, 8-24 May 2007 [ONLINE]. Available from: www.wild- cat.org/nigripes/infos/SliwaReportBenfSA07.pdf

159

SLIWA, A., WILSON, B., LAMBERSKI, N. & HERRICK, J. 2008. Report on surveying and catching Black-footed Cats (Felis nigripes) on Benfontein Game

Farm, 21-30 April 2008 [ONLINE]. Available from: http://www.wild- cat.org/nigripes/infos/Sliwa-et-al-report08.pdf

SLIWA, A., WILSON, B., LAMBERSKI, N., LAWRENZ, A. 2009a. Report on surveying and catching Black-footed cats (Felis nigripes) on Benfontein Nature

Reserve / Nuwejaarsfontein, 2–20 February 2009 [ONLINE]. Available from: www.wild-cat.org/nigripes/infos/Sliwa-et-al-report09.pdf

SLIWA, A., WILSON, B., LAMBERSKI, N. & LAWRENZ ,A. 2009b. Report on surveying and catching Black-footed Cats (Felis nigripes) on Benfontein Game

Farm / Nuwejaarsfontein, 9-19 November 2009 [ONLINE]. Available from: http://www.wild-cat.org/nigripes/infos/Sliwa-et-al2009-

Nuwejaarsfontein+Benfontein-report.pdf

SLIWA, A., WILSON, B., LAMBERSKI, N. & LAWRENZ, A. 2010. Report on surveying and catching Black-footed Cats (Felis nigripes) on

Nuwejaarsfontein/Benfontein Nature Reserve, 4-20 July 2010 [ONLINE].

Available from: http://www.wild-cat.org/nigripes/infos/Sliwa+Wilson+Lawrenz2010-

Report-Felis-nigripes-SA.pdf

160

SLIWA, A., WILSON, B., LAMBERSKI, N., LAWRENZ, A. & HERRICK, J. 2011.

Report on surveying, catching and monitoring black-footed cats (Felis nigripes) on

Nuwejaarsfontein Farm/Benfontein Nature Reserve in 2011 [ONLINE]. Available from: www.wild-cat.org/nigripes/infos/Sliwa+al2013-Report-Felis-nigripes-

SA2012.pdf

SLIWA, A., WILSON, B., LAMBERSKI, N., LAWRENZ, A. 2013. Report on surveying, catching and monitoring Black-footed cats (Felis nigripes) on

Benfontein Nature Reserve, Nuwejaarsfontein Farm, and Biesiesfontein in 2012

[ONLINE]. Available from: www.wild-cat.org/nigripes/infos/Sliwa+al2013-Report-

Felis-nigripes-SA2012.pdf

SLIWA, A., WILSON, B., LAMBERSKI, N. & TORDIFFE, A. 2014. Report on surveying, catching and monitoring black-footed cats (Felis nigripes) on Benfontein

Nature Reserve, Nuwejaarsfontein and Taaiboschpoort Farms in 2013 [ONLINE].

Available from: www.wild-cat.org/nigripes/infos/Sliwa+al2014-Report-Felis- nigripes-SA2013.pdf

SLIWA, A., WILSON, B., KUSTERS, M., LAWRENZ, A., EGGERS, B., HERRICK,

J., TORDIFFE, A., MARAIS, P. & MARAIS, S, A. 2015. Report on surveying, catching and monitoring black-footed cats (Felis nigripes) on Benfontein Nature

Reserve, Nuwejaarsfontein and Taaiboschpoort Farms in 2014 [ONLINE].

Available from: www.wild-cat.org/nigripes/infos/Sliwa_al2015-Report-Felis- nigripes-Work-SA2014.pdf

161

SMITHERS, R.H.M. 1971. The mammals of Botswana. Museum Memoir No. 4.

Salisbury: The Trustees of the national Museums of Rhodesia.

SMITHERS, R.H.M. 1983. The mammals of the southern African subregion.

Pretoria: University of Pretoria.

SMITHERS, R.H.M. 1986. South African Red Data Book – terrestrial mammals.

Foundation for Research Development, South Africa.

SOUTH AFRICA. 2004. The National Environmental Management Biodiversity

Act (NEMBA) No. 10. Government Gazette 487(26426), 7 June 2004.

STUART, C.T. 1982. The distribution of the small-spotted cat, Felis nigripes. The

Naturalist 26(3):8-9.

STUART, C.T. & STUART, T. 1988. Field guide to the mammals of southern

Africa. Cape Town: Struik Publishers.

STUART, C.T. & WILSON, V.J. 1988. The cats of southern Africa. Bulawayo: The

Chipangali Wildlife Trust.

SUNQUIST, M. & SUNQUIST, F. 2002. Wild cats of the world. London:

University Chicago Press.

162

TERIO, K.A., O’BRIEN, T., LAMBERSKI, N., FAMULA, T.R., & MUNSON, L.

2008. Amyloidosis in black-footed cats (Felis nigripes). Veterinary

Pathology 45(3):393-400.

UNITED NATIONS ENVIRONMENT PLAN (UNEP). 2002. Africa Environment

Outlook – past, present and future perspectives. Nairobi: Division of Early Warning and Assessment (DEWA), United Nations Environment Programme (UNEP).

[Online]. Available from: www.unep.org/dewa/africa/publications/aeo-1/index.htm

UNITED NATIONS ENVIRONMENT PLAN (UNEP). 2008. Africa: Atlas of our changing environment. Nairobi: Division of Early Warning and Assessment

(DEWA), United Nations Environment Programme (UNEP).

VAN DER MERWE, P., SAAYMAN, M. & ROSSOUW, R. 2014. The economic impact of hunting: a regional approach. South African Journal of Economic and

Management Sciences 17(4):379-395.

VISSER, J. 1976. Status and conservation of the smaller cats of southern Africa.

In: Eaton, R.L. (ed.). Contributions to status, management and conservation:

Proceedings of the Third International Symposium on the World’s Cats, 26-28 April

1974. Seattle: University of Washington:70.

VOLLMAR, A., MACKLIN, J.A. & FORD, L.S. 2010. Natural history specimen digitization: challenges and concerns. Biodiversity Informatics 7:93-112.

163

WATSON, J.P. 2006. Check list of the mammals of Tussen-die-Riviere Provincial

Nature Reserve, Free State Province, South Africa. Koedoe 49(1):111-117.

WEBSTER, C.M. & DESTEFANO, S. 2004. Using public surveys to determine the distribution of greater roadrunners in urban and suburban Tucson, Arizona. In:

Shaw, W., Harris, L.K., van Druff, L. (eds.). Proceedings 4th International Urban

Wildlife Symposium. Tucson, AZ: University of Arizona:69-72.

WERDELIN, L., YAMAGUCHI, N., JOHNSON, W.E. & O’BRIEN, S.J. 2010.

Phylogeny and evolution of cats (Felidae). In: Macdonald, D.W. & Loveridge, A.J.

(eds.). Biology and Conservation of Wild Felids. Oxford: University Press:59-82.

WILSON, B & SLIWA, A. 2007. Black-footed Cat Felis nigripes geographic range.

In: Sliwa, A. 2008. Felis nigripes. The IUCN Red List of Threatened Species.

Version 2014.2. [Online]. Available from: www.iucnredlist.org [Accessed:

15/09/2014].

WILSON, B., SLIWA, A. & DROUILLY, M. in prep. A Conservation Assessment of Felis nigripes. In: Child, M.F, Roxburgh, L. Do Linh San, E., Raimondo, D.

Davies-Mostert, H.T. (ed.). The Red List of Mammals of South Africa, Swaziland and Lesotho. South African National Biodiversity Institute and Endangered Wildlife

Trust, South Africa.

164

WOZENCRAFT, W.C. 1993. Order Carnivora. In: Wilson, D.E. & Reeder, D.M.

(eds.). Mammal species of the world, a taxonomic and geographic reference vol.

1. Washington DC: Smithsonian Institution Press:536.

WYNBERG, R. 2002. A decade of biodiversity conservation and use in South

Africa: tracking progress from Rio Earth Summit to the Johannesburg World

Summit on Sustainable Development. South African Journal of Science 98

(May/June):233-243.

ZAMBIA. Ministry of Tourism, Environment and Natural Resources, 2009. United

Nations Convention on Biological Diversity Fourth National Report.

ZIMBABWE. Ministry of Environment & Natural Resources Management,

December 2010. Zimbabwe’s Fourth National Report to the Convention on

Biological Biodiversity.

ZIMMERMANN, P.A., LAWRENZ, A. & SLIWA, A. 2011. Untersuchungen zu

Amyloidose und Akute-Phase- Proteinen bei Schwarzfußkatzen (Felis nigripes).

Tierärztl. Umschau 66, 364 – 368.

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ANNEXURES

ANNEXURE A: PUBLIC AWARENESS POSTER: ENGLISH

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ANNEXURE B: PUBLIC AWARENESS POSTER: AFRIKAANS

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ANNEXURE C: QUESTIONNAIRE: COMPLETED EXAMPLE

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ANNEXURE D: ETHICAL CLEARANCE

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