1 Caracals in a Heterogeneous Landscape

Caracals in a Heterogeneous Landscape: Resolutions for Human-Carnivore Conflicts

Item Type text; Electronic Dissertation

Authors Neils, Aletris Marie

Publisher The University of Arizona.

Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction or presentation (such as public display or performance) of protected items is prohibited except with permission of the author.

Download date 26/09/2021 18:05:44

Link to Item http://hdl.handle.net/10150/627732

CARACALS IN A HETEROGENEOUS LANDSCAPE: RESOLUTIONS FOR HUMAN-CARNIVORE CONFLICTS

Aletris Marie Neils

A Dissertation Submitted to the Faculty of the

SCHOOL OF NATURAL RESOURCES AND THE ENVIRONMENT

In Partial Fulfillment of the Requirements

For the Degree of

DOCTOR OF PHILOSOPHY

WITH A MAJOR IN NATURAL RESOURCES

In the Graduate College

THE UNIVERSITY OF ARIZONA

2018

STATEMENT BY AUTHOR

This dissertation has been submitted in partial fulfillment of the requirements for an advanced degree at the University of Arizona and is deposited in the University Library to be made available to borrowers under rules of the Library.

Brief quotations from this dissertation are allowable without special permission, provided that an accurate acknowledgement of the source is made. Requests for permission for extended quotation from or reproduction of this manuscript in whole or in part may be granted by the head of the major department or the Dean of the Graduate College when in his or her judgment the proposed use of the material is in the interests of scholarship. In all other instances, however, permission must be obtained from the author.

SIGNED: Aletris Marie Neils

ACKNOWLEDGEMENTS

I would like to begin by expressing my gratitude to my local farming hosts at Kiripotib

Guest Farm, Hans and Claudia von Hase, your lovely family and the whole Kiripotib team! You gave me my Namibian home away from home. I owe you so much- keep farming grass! Many thanks to Tyse and Annetta Swart for being the first family to allow me to release a radio collared caracal back onto their farm, for teaching me to “get out of my western way of seeing wildlife conflicts”, and for hours of great discussions and Afrikaans lessons. Thank you Timm and Inez

Miller- your incredible farm embodies what conservation farming can and should be. You are the future of Namibian farming and personify that coexistence is not just a dream! I also thank Udo and Tenda Bartsch for all your support. Thank you, Pierre and Select Cars, for keeping my bakkie

(truck) running after the thousands of hard kilometers I drove, I literally would have been stuck without you. I also acknowledge the hundreds of livestock farmers that contributed specimens, assistance, information, stories, and cups of rooibos tea. I could not have done it without your cooperation and trust.

Field research was conducted with meticulous detail in part by the exceptional Elin

Gromberg and Magnus Nystrand- thank you both for doing an incredible job continuing this research while I was back in the U.S. I am very grateful for field assistance from several passionate interns especially my amazing little sister (and fledgling conservationist) Elan Prince Henry Fox,

Lily Harrison, Amanda Veals, Colleen Cavanaugh, Allie Pena, and Sean Thomann and many others who contributed to the success of the project through the years. Necropsy protocols were developed with direction and guidance from Dr. Axel Hartmann, Lizzy Hartmann, and the

Otijiwarongo Veterinary Clinic – thank you for devoting so much time to training me. I also thank 4

the Bitterwasser Flying Center for accomidations and support. I am appreciative for Southwest

Wildlife Conservation Center, particularly Linda Searles and for allowing me to refine collars and methodology before deploying them in the field. This research was conducted with support from the Namibian Ministry of Environment and Tourism (under permit number 1539). The American

Cultural Center in Windhoek was very helpful in assisting me in country. Funding was provided by Fulbright, Cheetah Kids, Crosby Family, Bonnie Kay, Martha Sloan, and contributions from many other private donors – without you support these caracals would never have been able to be released.

I thank my advisors Dr. John Koprowski and Dr. Cecil Schwalbe for all of their assistance throughout this project, and for granting me complete freedom but being there when I needed help most. John without your patience, motivation, and guidance in navigating this process I would not be here. Cecil, there exists no better cheerleader! Thank you for your endless enthusiasm. I also greatly appreciate Dr. Tom Sheridan for encouraging me to explore the social science component of this work and guiding me though the complexities of the “soft science”- this turned out to be the most important piece to the puzzle and I would not have discovered it without you. I would also like to thank Dr. Kevin Bonine for his ecological insight, support, and for being a mentor on disseminating science to the public. Thank you, William Shaw, for your insight on the social dimensions of wildlife managements. Thank you all for your guidance and constructive comments.

I thank Diane Austin for her advice and direction in my qualitative interview templates and for teaching me the nuances of social science methodology. I thank Cheyenne Rose McChesney for her assistance transcribing and coding. I owe a debt of graditute to Katie Hughes (SNRE) and

Lori D’Anna (Graduate College) for technical support and helping navigate hurdles and make 5

deadlines. Thank you, ladies, for everything! I would also like to thank my eighth-grade science teacher Mrs. Eileen Jones for transforming my views of science class and inspiring me to keep my career trajectory on “scientist”.

I would also like to thank the many mentors who have helped me get to where I am today, especially the legendary David E. Brown, who since I was a teenager has showed me how to be a naturalist and holistically understand the landscape I am in. The encyclopedic ecologist, Randy

Babb was always willing and helpful to “talk Africa” with. Thank you Dr. Andrew Smith for opening my eyes to the amazing diversity of mammals. I am grateful to Dr. Laurie Marker and the Cheetah Conservation Fund for being my platform while I was getting started and being my introduction to human-carnivore conflict in Namibia. Without Jack Childs introducing me to apex cats (nearly twenty years ago now), I never would have ventured to Namibia in the first place – for this and for all you taught me about how to track predators I am eternally grateful.

I am also indebted to the dynamic Sandy Anderson for her inspiration and helping me cut through the crap and focus on what matters – you are the architype that number of degrees should not be the rubric on knowledge. I am so grateful to Andy Salazar for showing me how to trail with hounds and for sharing all his insights on predators – you will always be remembered. Thank you to the late William ‘Sky’ Crosby for your long conversations, great questions, infectious laugh, and all you have done for conservation across the globe.

I am so blessed to have found such great friendship with Bella Hilda – thank you for showing me how Namibian farmers live and for all the great picnics with Stephan, Klara, and

Dawid in the veld. To Elizabeth ‘Lizard’ Ridder thank you for your friendship, love, and support.

To my best friend Paige McNickle, thank you for your great advice, for always being a shoulder I can lean on, and for always believing in me. To my best amigo Matt Skroch, thank you for your 6

support, long conversations, direction, and of course for introducing me to the sky islands. Julia

Muldoon Marquez, thank you for your friendship and for loving our critters while we were traveling – I can’t wait to see the great contributions you make in this field.

I am forever grateful to the Carver family – John, Jody, Cameron and Kristin for all of your support getting started. Without your organization, CheetahKids, I would have never been able to radio-collar the first caracals. Your family is so special to me.

Finally, I would like to thank my family for always supporting me and sending their love and care packages while I was overseas for extended periods of time. I am so privileged to have been born with parents that fostered my love for nature, encouraged me to explore outside, and tolerated all the specimens I was always collecting. Without my huge, dynamic, loving family

(Michael Neils, Kathleen Henry, Yaro Shon Neils, Raemi Neils, Courtney (and Winston) Neils,

Elan Fox, Betty Neils, Liam Neils, Eliza Neils, Stephanie, Heath & Ben) I could not have accomplished this monumental task- it would not have been possible. My grandma Betty gave me my first book on African Cats when I was five years old and is responsible for planting the seed for this work; thank you for watching over me! I am also grateful for my late uncle Brett for always being so incredibly proud of us and for always wanting to hear my stories about Africa; I hope you can see it now. My grandpa Bill shared his love for adventure and cats, and my grandma

Marie shared her admiration for African wildlife, and my uncle Benny (Benton Henry) helped shape how I see things. I am also so appreciative of my in-laws Rebecca and Ted Crosby and Don

Bugbee and my Connecticut family for your love, encouragement, and support (and for raising such a great son). Thank you to my parents for encouraging me to dream big, and always reach for my dreams. Many thanks to my mom Kathleen Henry and sister Yaro Neils for taking care of our house and caring for our critters so Chris could join me overseas for periods of time. I thank 7

my clan of remarkable sisters for all their love, support and encouragement, and even more love – always and unconditionaly!

Most notably, I thank my husband and partner, Chris Bugbee. Thank you for your contribution every step of the way; you deserve a PhD every bit as much as me. You are my rock that has gotten me though the many, many, many hurdles I encountered throughout this project

(and life during this time). I could not have done this without you – period. Since we made it through this, I know we can make it through anything and everything. I am most grateful for our beautiful, precious Aurelia- you are my life’s greatest achievement and after labor made this dissertation seem easy. Your laugh and smile fueled us to the finishline.

DEDICATION

I have a double dedication: I must dedicate this work to Namibia, to both its incredible wildlife and people – it remains one of the last wild places on the planet and I will forever appreciate how wild Namibia changed my life. I also dedicate this dissertation to Leopold (Nubs) Neils – you were a majestic cat who inspired me to devote my life to saving your wild cousins. May I make you proud!

TABLE OF CONTENTS

LIST OF TABLES ...... 14

LIST OF FIGURES ...... 16

ABSTRACT ...... 18

INTRODUCTION...... 20

PRESENT STUDY...... 22

LITERATURE CITED ...... 23

CARACAL CARACAL (Carnivora: Felidae) ...... 24

Phylogenetic Relationship- ...... 24

Physical Description- ...... 25

Distribution- ...... 27

Habitat- ...... 27

Diet and Feeding Ecology- ...... 28

Predators- ...... 32

Behavior and Social Structure- ...... 33

Reproduction- ...... 34

Conservation Status- ...... 35

Literature Cited ...... 39

Figures ...... 44

APPENDIX A: DIET OF CARACALS (CARACAL CARACAL) ON NAMIBIAN

FARMLANDS ...... 45

Diet of Caracals (Caracal caracal) on Namibian Farmlands ...... 46 10

METHODS ...... 49

RESULTS ...... 55

DISCUSSION ...... 59

ACKNOWLEDGMENTS ...... 66

LITERATURE CITED ...... 67

APPENDIX A ...... 73

FIGURE LEGENDS ...... 77

TABLES ...... 78

FIGURES ...... 85

APPENDIX B: SURVIVAL, MORTALITY & REPRODUCTION OF CARACALS

(CARACAL CARACAL) ON NAMIBIAN FARMLANDS ...... 86

INTRODUCTION ...... 89

METHODS ...... 91

2.1. Study area ...... 91

2.2. Animal trapping, handling, and immobilization ...... 92

2.3. Monitoring ...... 93

2.4. Survival and reproduction ...... 94

2.5. Life tables ...... 94

2.6. Incorporating estimated age and reproduction into life tables ...... 96

2.7. Survival curves ...... 96

RESULTS ...... 97

3.1. Captures and necropsies ...... 97

3.2. Life tables ...... 98 11

3.4. Survival curves ...... 98

3.5. Reproduction ...... 99

DISCUSSION ...... 99

4.1. Felid response to high mortality ...... 100

4.2. Population sinks ...... 101

4.3. Caracal kitten sex ratios and survival ...... 102

4.4. Long-term ecological consequences ...... 103

4.5. Conservation implications ...... 104

LITERATURE CITED ...... 107

TABLES ...... 113

FIGURE LEGENDS ...... 119

FIGURES ...... 120

APPENDIX C: UNINTENDED CONSEQUENCES: HOW THE ANIMAL RIGHTS

MOVEMENT INADVERTENTLY LED TO THE PERSECUTION OF NAMIBIAN

CARNIVORES ...... 123

INTRODUCTION ...... 125

METHODS ...... 126

2.1. Study site ...... 126

2.2. Subjects ...... 127

2.3 Interviews ...... 127

2.4 Analysis ...... 129

RESULTS ...... 129

3.1. Farm characteristics-...... 129 12

3.2. Livestock and predation-...... 130

DISCUSSION ...... 133

4.1. Predator perceptions and persecution- ...... 133

4.2. Effects of culling on carnivore populations ...... 134

4.3. Legal ownership of wildlife- ...... 136

4.4. Drought and climate change- ...... 137

4.5. Livestock breeds- ...... 137

4.6. The Anti-Fur Movement’s impact to Namibian farmers- ...... 141

4.7. Conservation implications of market shift- ...... 143

4.8. Conclusions ...... 147

ACKNOWLEDGEMENTS ...... 149

REFERENCES ...... 150

TABLES ...... 155

FIGURES ...... 157

APPENDIX D: Permits ...... 158

APPENDIX E: Caracal Necropsy Form ...... 162

APPENDIX F: Farmer Qualitative Interview Template ...... 164

APPENDIX G: Capture Checklist ...... 166

APPENDIX H: Curriculum Vitae ...... 166

VITA...... 179

LIST OF TABLES

Table A1. Total prey items in the diets of southern Namibian caracals (Caracal caracal) as determined by kill site, stomach content and scat analyses from 2010-2016...... 78

Table A2. Relative frequencies of prey items in the diets of southern Namibian caracals

(Caracal caracal) taken from Karoo/Karasberg (KK), Kalahari grassland (KG), sparse shrubland (SS) and Acacia thornbrush (AT) habitats from 2010-2016...... 79

Table A3. Relative frequencies of prey items in the diets of southern Namibian caracals

(Caracal caracal) represented by three different methods including stomach contents, kill sites and scat samples from 2010-2016...... 80

Table A4. Relative frequencies of prey items in the diets of southern Namibian caracals

(Caracal caracal) in Karoo/ Karasberg habitat as represented by three different methods including stomach contents, kill sites and scat samples from 2010-2016...... 81

Table A5. Relative frequencies of prey items in the diets of southern Namibian caracals

(Caracal caracal) in Kalahari grassland habitat as represented by three different methods including stomach contents, kill sites and scat samples from 2010-2016...... 82

Table A6. Relative frequencies of prey items in the diets of southern Namibian caracals

(Caracal caracal) in sparse shrubland habitat as represented by three different methods including stomach contents, kill sites and scat samples from 2010-2016...... 83

Table A7. Relative frequencies of prey items in the diets of southern Namibian caracals

(Caracal caracal) in Acacia thornbrush habitat as represented by three different methods including stomach contents, kill sites and scat samples from 2010-2016...... 84

Table B1. Numbers of yearling/ sub-adult (Y/ SA) to adult (A) caracals (Caracal caracal) on southern Namibian farmlands monitored from 2010-2016...... 113

Table B2. Life table statistics calculated for all caracals (Caracal caracal) of known age from southern Namibian farmlands collected from 2010-2016...... 114

Table B3. Life table statistics calculated for all male caracals (Caracal caracal) of known age from southern Namibian farmlands collected from 2010-2016...... 115

Table B4. Life table statistics calculated for all female caracals (Caracal caracal) of known age from southern Namibian farmlands collected from 2010-2016...... 116

Table B5. Life table statistics calculated for all female caracals (Caracal caracal) of known age from southern Namibian farmlands collected from 2010-2016...... 117

Table B6. Kaplan-Meier survival curve statistics (in days) for male, female, yearling/ subadult, and adult radio-collared caracals (Caracal caracal) from three different habitat types (Karoo/ Karasberg, Khalahari grassland; sparse shrubland) on southern Namibian farmlands monitored from 2010-2016...... 118

Table C1. Characteristics of small stock found on Namibian farmlands...... 155

Table C2. Minimum population estimates of livestock by breed by year in Namibia...... 156

LIST OF FIGURES

Figure 1. Map of current geographic range of Caracal caracal based on available data

(2018)…...... 43

Figure A1. Study area for southern Namibian caracal (Caracal caracal) dietary study with habitat delineations for Karoo/Karasberg, Kalahari grassland, sparse shrubland and Acacia thornbrush habitats...... 85

Figure B1. Kaplan-Meier survival curve for male and female radio-collared caracals (Caracal caracal) from southern Namibian farmlands monitored from 2010-2016 (P = 0.0573)...... 120

Figure B2. Kaplan-Meier survival curve for yearling/ subadult (Y/ SA) and adult (A) radio- collared caracals (Caracal caracal) from southern Namibian farmlands monitored from 2010-

2016 (P = 0.0002)...... 121

Figure B3. Kaplan-Meier survival curve for radio-collared caracals (Caracal caracal) from three different habitat types (Karoo/ Karasberg, KK; Kalahari grassland, KG; sparse shrubland,

SS) on southern Namibian farmlands monitored from 2010-2016 (P = 0.9615)...... 122

Figure C1. Model depicting the chain of events that followed the international anti-fur movement which led to the decline of pelt sheep in Namibia and an increase of predators removed. Data were collected via semi-structured interviews...... 157

Figure D1. Initial research and collection permit issued by the Namibian Ministry of

Environment and Tourism for the caracal research project...... 159

Figure D2. Initial research and collection permit issued by the University of Arizona

Institutional Animal Care and Use Committee for the caracal research project...... 160

Figure D3. Social science research protocal by the University of Arizona’s Human Subjects

Protection Program...... 161

ABSTRACT

In regions where carnivores and livestock coexist, conflicts are virtually inevitable.

Ecological, economic and social consequences of the interaction between predator and prey exist, especially in places where predators and stock producers share the same land and resources. Throughout southern Africa, caracals are regarded as vermin due to occasional predation on livestock. Goals of this research were to examine specific life history traits that render caracals susceptible to conflict with humans, to understand impacts of human persecution on caracal ecology, and to offer alternative strategies for coexistence that work within the context of caracal natural history. Data were collected from 34 radio-collared and 208 necropsied caracals on working farms throughout Namibia from 2010 to 2016. Relative frequencies of prey items in caracal diet were calculated from analyses of 202 caracal stomachs,

688 kill sites and 250 caracal scats (N=1140 samples). A total of 106 species were recorded in

Namibian caracal diet. Mammalian prey made up 83.2% of diet, however, stock comprised only

2.1% of caracal diet despite their abundance throughout the study areas. Regardless of relatively insignificant depredation rates, perception of caracals as livestock killers results in extensive persecution by farmers. Demographic parameters of caracals, including survival and reproduction values, were estimated and used to generate life history schedules and Kaplan-

Meier survival curves to determine whether caracal populations on southern Namibian farmlands can withstand high anthropogenic mortality. Life table data revealed an estimated net reproductive rate of 0.891- 1.06. Southern Namibia stock farms likely represent a mosaic of population sinks for caracals where populations rely on immigration from adjacent areas to

persist; demography of caracal populations on farmlands is highly skewed towards young male dispersers. Data indicate that caracal populations are projected to decline if extensive persecution pressure continues. The underlying causes of this persecution was investigated, using 561 qualitative interviews from 367 Namibian farmers, to enhance understanding of wildlife conflicts and risk perception from a socio-economic context with the goal of identifying management actions to reduce the number of predators killed by people. Respondents indicated that drivers of conflict include drought, resource availability, and market-driven demands for certain breeds of sheep whose husbandry practices lead to increased conflicts with predators.

Results suggest farming systems exist in Namibia where predator eradication is the primary strategy over livestock management. Consequences of this extensive conflict are not sustainable for stock producers or caracals in the long-term. Conclusions of this project have implications for carnivore conflicts in parallel milieus worldwide.

INTRODUCTION

Caracals (Caracal caracal) are a poorly understood wild felid that have garnered relatively little research attention. Few studies have been published of their basic ecology, behavior, and life history traits and sample sizes have been small (Marker & Dickman 2005; Sunquist & Sunquist

2002). No science-based population estimates exist for caracals even though caracals are considered threatened, endangered or on the verge of extinction throughout a majority of their range (Sunquist & Sunquist 2002). In sub-Saharan Africa, caracals are protected in about half of their range states (Nowell & Jackson 1996). Throughout most of southern Africa, the last remaining geographic stronghold for caracals throughout their range, caracals receive no legal protection and are highly persecuted as vermin due to occasional predation on small stock

(Avenant & Nel 1998). Namibian farmers consider caracals the top threat to small stock and the fourth greatest threat to their cattle (Stein 2006). Landowners may lethally remove them without restriction.

Prior to the 1990s, most research programs on caracals consisted of finding effective means of exterminating the species. From 1931-1952 over 2,200 caracals per year were killed in control operations in the Karoo, South Africa (Stuart 1984). Similarly, farmers in Namibia reported killing up to 2,800 caracals in a single year and much killing likely goes unreported (Nowell & Jackson

1996). Records on numbers of caracals killed each year are probably gross underestimations since caracal pelts are of little commercial value (Sunquist & Sunquist 2002). Although data are limited, persecution has lasted for decades on a regional scale and continues to this day (Stuart & Stuart

2013). Caracals have appeared to so far withstand rigorous eradication campaigns in South Africa

and Namibia, but numbers may be dwindling. Population assessments are needed to determine if declines have occurred.

The objectives of this study were threefold. The first was to document caracal life history traits and mortality rates on Namibian farmlands. Conservation and management of species experiencing heavy anthropogenic impacts can greatly benefit from detailed demographic information; future population collapse can be predicted by examining population demographics and age distributions. By examining vital statistics of caracals on Namibian farmlands, status and population trends of caracals can be assessed. Such data have never been collected for caracals.

The second objective was to examine caracal diet on southern Namibian farmlands to determine the extent of livestock depredation and what aspects of their environment, biology or ecology predispose them to depredation. The threat caracals pose to domestic livestock could be less than commonly perceived, as carnivores on the landscape are a common scapegoat for suboptimal stock production and unexpected stock losses and death (Kellert et al. 1996, Treves & Karanth 2003,

Musiani & Paquet 2004). Scientific examination of livestock conflict in southern Namibia requires thorough understanding of caracal natural history, dietary habits, and whether predation varies by habitat. The third objective was to better understand farmer relationships with and perceptions of caracals and other predators on the landscape. From a socio-political perspective, extensive qualitative interviews with local commercial farmers were conducted to understand the nuances of conflict and recognize effects depredation has on livelihoods. Scientific research cannot translate into conservation action without addressing all sides of the conflict.

PRESENT STUDY

My dissertation includes an introduction and natural history of Caracal caracal and three manuscripts formatted for submission to peer-reviewed journals. The first manuscript, intended for submission to the Journal of Mammalogy, (Appendix A), “Diet of Caracals (Caracal caracal) on Namibian Farmlands” documents results from a comprehensive caracal diet study using scat, stomach content, and killsite analyses in four distinct ecosystems in southern Namibia. The second manuscript, intended for submission to Biological Conservation, (Appendix B), “Survival, mortality and reproduction of caracals (Caracal caracal) on Namibian farmlands” documents caracal survival and reproduction rates used to generate life history schedules and Kaplan-Meier survival curves for the southern Namibian population. The third manuscript, intended for submission to Human Ecology, (Appendix C), “Unintended consequences: How the animal rights movement inadvertently led to the persecution of Namibian carnivores” documents Namibian farmers’ perceptions of the risk posed by predators to their livelihoods using semi-structured interviews. Methods, results, and conclusions for this study are presented in the appended manuscripts.

LITERATURE CITED

Avenant, NL, Nel, JAJ. 1998. Home-range use, activity, and density of caracal in relation to prey density. African Journal of Ecology 36:347-359.

Kellert, SR, Black, M, Rush, CR, Bath, A. 1996. Human culture and large carnivore conservation in North America. Conservation Biology 10:977-990.

Marker, L, Dickman, M. 2005. Notes on the spatial ecology of caracals (Felis caracal) with particular reference to Namibian farmlands. African Journal of Ecology 43:73-76.

Musiani, M, Paquet, PC. 2004. The practices of wolf persecution, protection, and restoration in Canada and the United States. BioScience 54:50-60.

Nowell, K, Jackson, P. 1996. Wild cats: status survey and conservation action plan. Gland: IUCN, 382.

Stein, A. 2006. Interactions: Leopards vs. Brown Hyenas. Otjiwarongo, Namibia: Waterberg Carnivore Project Namibia.

Stuart, CT. 1984. The distribution and status of Felis caracal Schreber, 1776. Saugetierkundliche Mitteilungen. 31:197- 203.

Stuart, C, Stuart, T. 2013. Caracal Caracal caracal: Mammals of Africa 5:174-179.

Sunquist, M, Sunquist, F. 2002. Wild cats of the World. The University of Chicago Press, Chicago.

Treves, A, Karanth, KU. 2003. Human‐carnivore conflict and perspectives on carnivore management worldwide. Conservation Biology 17:1491-1499.

CARACAL CARACAL (CARNIVORA: FELIDAE)

Caracals (Caracal caracal) are among the least studied felids in the world with little known of their ecology, behavior and population status (Singh et al. 2014). Caracals are rare, endangered or extirpated in the northern half of their range (Sunquist & Sunquist 2002). In southern Africa, however, caracals are classified as ‘vermin’ and may be destroyed without restriction. Prior to the 1990s, most research programs on caracals focused on finding effective means of exterminating the species (Stuart & Stuart 2013). No science-based population estimates exist for caracals in any part of their range. Considering high rates of direct anthropogenic mortality in the species’ geographic stronghold, caracals are probably the most persecuted felid in the world.

Phylogenetic Relationship-

The closest genetic relatives to caracals are African golden cats (Profelis aurata) and servals (Leptailurus serval) (Johnson et al. 2006). These three species form the Caracal clade, a lineage that diverged from other extant felids approximately 9.4 million years ago (Johnson et al.

2006). Captive hybrids have led some authorities to classify servals in the Caracal genus

(Johnson et al. 2006). Nine subspecies have been described although genetic profiles are lacking. The southernmost two subspecies (C. c. caracal and C. c. damarensis) are the only ones that have garnered any significant research attention (Stuart 1986).

Physical Description-

Caracals are a long, slender, medium-sized cat. Head and body lengths range from 610-

1057 mm (average 692-881 mm) with the largest individuals from South Africa and the smallest from Israel (Stuart 1981; Smithers 1971; Gaillard 1969; Nader 1984; Sunquist & Sunquist 2002).

Caracals stand from 40-50 cm at shoulder height with tail lengths ranging from 100-340 mm

(Smithers 1983). Compared to most cats, caracals have a relatively short tail, roughly one third of the body length. Weights range from 5.8-20 kg, with a range-wide average of 12.7 kg for males and 10.1 kg for females (Roberts 1977; Pringle & Pringle 1979; Heptner & Sludski, 1992;

Stuart 1981; Smithers 1971; Gaillard 1969; Nader 1984; Smithers & Wilson 1979; Smithers

1983; Weisbein & Mendelssohn 1990; Sunquist & Sunquist 2002).

Caracal pelage is nearly uniform in color and may be tan, brown, grayish or reddish, often approximating hues of local or regional geological features (Smithers 1983). Pelage is lighter and spotted on the throat, chin, belly and underside of limbs (Smithers 1983).

Intraspecific color variations are slight but female caracals are generally lighter in color

(Sunquist & Sunquist 2002). Melanistic individuals have been recorded from central Africa in tropical habitat (Happold 1987). Caracal fur is short, dense and provides effective insulation from extreme temperatures that characterize much of the species’ range (Smithers 1983). Guard hairs are 15-30 mm with light bases and grizzled silver, gray or black tips. Seasonal variation occurs in coat thickness and degree of underfur (Pocock 1939).

An immediately recognizable feature of caracals are tall triangular ears with long black ear tufts. Although some other felids exhibit ear tufts, notably members of the Lynx genus, caracal tufts are the most extreme. Tufts may grow longer with age; in older animals tufts hang 25

down like tassels (Kingdon 1977). Ear tufts may serve as disruptive camouflage, breaking up the outline of the caracal in tall, swaying grasses. Like satellite dishes, caracal ears work independently from one another to hone on target soundwaves from up to 200m away (Smithers

1983).

Caracal hind legs are longer than front legs, giving an overall sloped appearance.

Powerfully built hindquarters enable caracals to execute explosive bursts of speed and vertical leaps >3m in the air to acquire avian prey. Caracal muscle fibers can generate 3x the power of human counterparts (Kohn & Noakes 2013). Paws are relatively large with four digits on the hind feet and five on the front. The fifth digit is a heavily built dewclaw utilized as a weapon for hunting (Grobler 1981). Caracals have stiff guard hairs on the underside of the feet, more pronounced in individuals that inhabit sandy environments, which may be advantageous for moving through soft substrate (Heptner & Sludskii 1992). Foot hairs probably increase ability to silently stalk prey.

Among extant felids caracal skulls are described as “generalized” with relatively massive morphological features and a rounded, blunt rostrum (Sicuro 2011). Auditory bullae are large, indicating a keen sense of hearing (Smithers 1983). Sagittal crest is distinctive, providing extra surface area for jaw muscle attachment translating to powerful jaw strength. Mandible is thick and stout with a tall coronoid process that provides additional leverage for muscles involved in closing the jaw (Smithers 1983). Orbital sockets are relatively large and caracals are thought to have excellent vision. Teeth are relatively large and well-enameled (Smithers 1983). Upper canines measure to 20 mm and are separated by a diastema both in front and behind adjoining teeth, while the lower canines have only a posterior diastema (Pringle & Pringle 1979). Second 26

premolars are rarely present in most individuals. These features, along with highly reduced lower incisors, allow canines to penetrate prey to full depth (Pringle & Pringle 1979).

Distribution-

Caracals have one of the widest distributions of extant carnivores, historically found in

North Africa from Morocco east to Egypt, Sudan, Ethiopia, southern Somalia, northern Niger and throughout sub-Saharan Africa excluding extreme Sahara and Namib deserts (Lynch 1983).

Their current distribution, however, is somewhat reduced (Fig.1). Caracals are replaced by

African golden cats in dense tropical forests (Smithers 1983). Caracals are recorded from the

Sinai Peninsula, Jordan, Syria, southeastern turkey, Arabia, Kuwait, Iraq, Iran, Turkmenia

(southern USSR), Afghanistan, Pakistan, and into India (Smithers 1983) yet are sparsely distributed and considered rare in these countries (Sunquist & Sunquist 2002). In South Africa and Namibia, the stronghold of the species, caracals are still relatively common and are typically regarded as vermin (Sunquist & Sunquist 2002). Their historic range mirrors that of the cheetah

(Anyinonyx jubatus).

Habitat-

Caracals typically inhabit arid and semi-arid regions throughout their range (Sunquist &

Sunquist 2002). Caracals prefer open habitats and tolerate hotter and drier conditions than L. serval (Sunquist & Sunquist 2002). Habitat preference varies slightly by region. In southern

Africa caracals are found in arid shrub savannah, woodlands, open grassland, Acacia scrub, and mixed riverine habitats (Smithers 1971). In South Africa’s Cape Provence caracals inhabit scrub pine and riverine habitats and in Kenya they are found primarily in grasslands. North African

caracals inhabit semiarid areas around massif mountain foothills; in Israel they are found in hilly grasslands with Acacia; in Pakistan they are found in arid subtropical scrub forests and tropical thorn forests (Sunquist & Sunquist 2002). Caracals typically associate with edge habitats where grasslands and forests meet (Heptner & Sludskii 1992; Kingdon 1977; Roberts 1977; Sunquist &

Sunquist 2002; Weisbein & Mendelssohn 1990). De Smet (1989) accounts one set of caracal tracks in the Sahara desert, but suggested that they probably don’t venture too far from dune edges. Caracals utilize mountainous terrain but generally stay close to the foothills (Moolman

1986), however, caracals are claimed to inhabit mountainous regions in high elevation pine forests (De Smet 1989). Regardless of habitat type, caracals require or prefer some form of adequate vegetative or rocky cover. Though they will cross wide open grassland during nighttime hours, this habitat is not primarily utilized without some kind of clumped vegetative structure (Smithers 1971).

Diet and Feeding Ecology-

Caracals are opportunistic predators documented predating on a broad range of prey including rodents, lagomorphs, hyraxes, small to mid-sized ungulates, small carnivores and reptiles (Grobler 1981; Moolman 1984; Moolman 1986; Skinner 1979; Smithers 1971; Stuart &

Hickman 1991; Weisbein & Mendelssohn 1989; Funston et al. 2001; A. Neils, personal observation). Diet is primarily mammal-based but varies by region, habitat, or locale (Moolman

1986; A. Neils, personal observation). Caracal habitat use is positively correlated with density of mammalian prey (Avenant & Nel 1998) and caracal foraging is best described as mammal-based searches, prey that caracals and other cat species are phylogenetically adapted to capture (Kok &

Nel 2004). Mammalian prey ranges in size from small rodents to large antelope such as springbok

(Antidorcas marsupialis), with mass of prey items typically averaging 45% the mass of the caracal

(Kok & Nel 2004). Avian prey ranges from quail to ostrich (Struthio camelus) (Smithers 1971), and reptilian prey ranges from small lacertid lizards to large varanids (Sunquist & Sunquist 2002).

Examination of southern Namibian caracal stomach contents, kill sites and scats revealed a total of 106 species including artiodactyls (39.4%), hyraxes (12.5%), birds (13.3%), rodents

(12.2%), lagomorphs (8.9%), carnivores (5.9%) and herpetofauna (3.6%) (A. Neils, personal observation). Similar diversity has been recorded in other studies and is indicative of the generalist feeding behavior of Namibian caracals (Grobler 1981; Moolman 1986; Palmer & Fairall 1988;

Skinner 1979; Smithers 1983; Stuart & Hickman 1991; Weisbein & Mendelssohn 1990). In

Mountain Zebra National Park, caracals primarily chose rock hyrax and mountain reedbuck

(Redunca fulvorufula) with mammals contributing 93.8% of the total ingesta (Grobler 1981).

Caracal prey in Karoo National Park consisted of rodents (39%), artiodactyls (28%), hyrax (22%), lagomorphs (19%), arthropods (17%), and birds and carnivores (2% respectively) (Palmer &

Fairall 1988). In Kgalagadi Transfrontier Park, caracals consumed 16 of the total 37 species of suitable mammal prey with which they shared their range, and 60% of the total mammal material consumed came from rodents (primarily springhare, Pedetes capensis), followed by lagomorphs

(Melville et al. 2004). In the Cape region of South Africa, rodents account for >70% of caracal diet, ungulates (Braczkowski et al. 2012). In India, 84% of all caracal scats analyzed contained rodents, which averaged 58% of the biomass contained within the scat (Mukherjee et al. 2004). In

Israel, caracals consumed mammals (62%), birds (24%) and reptiles (6%) (Weisbein &

Mendelssohn 1990). This is the highest proportion of avian prey reported in the literature except in Acacia thornbrush habitat in north-central Namibia (31%) (A. Neils, personal observation).

Caracals are considered an important species for controlling rodent populations (Pringle

& Pringle 1979), but unlike most small cats, caracals commonly take prey >2-3 times their own mass, a trend most often observed in larger felids (Grobler 1981; Sunquist & Sunquist 2002; A.

Neils, personal observation). Caracals are sufficiently equipped to kill large ungulates as opposed to preying almost exclusively on small mammals like the closely related serval (Leptailurus serval). The relative importance of large prey species vs. small mammals seems to vary by region.

In some ecosystems larger carnivores are extirpated resulting in caracals becoming the dominant predators (Sunquist & Sunquist 2002; Stuart & Stuart 2013). Without larger predators, caracals may have the ability to expand their niche and behave more like a higher-tiered predator. In areas where caracals coexist with larger felids, they may exhibit a different realized niche focusing on smaller prey such as rodents and birds as opposed to larger ungulates.

Caracal consumption of other carnivores has been noted by several authors (Lloyd & Millar

1981; Stuart 1984; Palmer & Fairall 1988; Melville et al. 2004; Braczkowski et al. 2012; A. Neils, personal observation). Black-backed jackal (Canis mesomelas) and African wildcat (Felis silvestris lybica) were the most commonly encountered carnivores preyed upon by caracals in

Namibia, but ten other species were recorded as well (A. Neils, personal observation). Black- backed jackals are anecdotally said to increase in areas where caracals are heavily persecuted

(Stuart 1981).

In southern Namibia, less than 12% of radio-collared caracals predated on domestic stock, which accounted for only 2.1% of overall caracal diet (A. Neils, personal observation). In 30

a coastal South African park where natural prey numbers were diminished, 15% of caracal diet was ungulates (both native antelopes and domestic stock) (Braczkowski et al. 2012). Several thousand sheep and goats are killed annually by caracals in the Cape Province of South Africa

(Stuart & Hickman 1991). In the Cape Province, stomach contents of caracals obtained from animal control operations were examined and one third of those stomachs that contained prey contained livestock (Stuart & Hickman 1991). Interestingly, 37% of those caracal stomachs were completely empty, indicating that caracals were struggling to acquire food resources in that area. Many studies involving caracal diet have been conducted near livestock farms that had diminished natural prey (Du Plessis et al. 2015).

Felids select against livestock if wild ungulate prey is abundant (Ackerman 1984; Biswas

& Sankar 2002; Reddy et al. 2004; A. Neils, personal observation). Caracal stock predation correlates with low natural prey densities, drought, and cold weather (Palmer & Fairall 1988;

Melville et al. 2004; Melville & Bothma 2006). Severity of livestock depredation by caracals is dependent on availability of wild prey and stock husbandry techniques (Stuart & Stuart 2013).

Some caracals on livestock farms never prey on livestock (Moolman 1986; Palmer & Fairall

1988; A. Neils, personal observation). Livestock may serve as a last resort for caracals unable to obtain wild prey due to inexperience, old age, injury, or management practices that deplete natural prey.

Like most felids, caracals deliver a killing bite either to the back of the neck (for relatively small prey) or to the throat (for relatively larger prey) (Sunquist & Sunquist 2002).

Some larger prey may also be dispatched with repeated bites and lacerations to the nape of the neck (Pringle & Pringle 1979). Caracals kill avian prey with a sideways slap of the front paws 31

delivered with great speed and force (Smithers 1971). Prior to an ambush, caracals often stalk to within 5 m of prey before initiating an explosively powerful attack. Caracals will often begin by consuming the head on smaller prey items (Skinner 1979). For larger ungulate prey caracals will begin by consuming the hindquarters, followed by the shoulders, and leaving the head intact

(Grobler 1981). Viscera are usually left untouched. If a kill is too large to consume in one sitting, caracals typically cache the remains and return to feed until the carcass is finished

(Grobler 1981). Caracals may wander large distances from cached carcasses in between consecutive feedings (A. Neils, personal observation). Some caracals have been observed feeding a single time on a large kill before abandoning it (Pringle & Pringle 1979). In areas of high human persecution this behavior may be a strategy to avoid being killed (Grobler 1981;

Stuart & Hickman 1991). Like most felids, caracals typically avoid feeding on carrion, although this behavior has been observed (Skinner 1979; Grobler 1981; Mendelssohn 1989).

Predators-

Caracals occasionally fall prey to larger sympatric felids and spotted hyenas (Crocuta crocuta) (Pienaar 1969; personal observation). In Kruger National Park a caracal was observed being killed by a pride of seven lions (Panthera leo) (Pienaar 1969). Besides male conspecifics, caracal kittens have been documented being killed by a variety of predators including leopards

(Panthera pardus), cheetahs, hyenas, black-backed jackals (Canis mesomelas), honey badgers

(Mellivora capensis), and rock pythons (Python sebae). Black-backed jackal predation on caracal young may explain reported surges in caracal populations following extreme jackal removal efforts (Pringle & Pringle 1979).

Behavior and Social Structure-

Caracals are an elusive species best surveyed from spoor and camera traps (Singh et al.

2014), and to a lesser extent by spotlighting and using audio playbacks (Thorn et al. 2010). They may be active any time in a 24-hr period, but are primarily nocturnal and crepuscular (Singh et al. 2014). Caracal activity is influenced by temperature and sunrise/sunset times (Weisbein &

Mendelssohn 1990). In cool cloudy weather, caracals may exhibit more diurnal behavior

(Smithers 1971).

Generally solitary, caracals are rarely observed in pairs. Such instances usually involve a female with older offspring, but occasionally breeding adults will travel together temporarily

(Smithers 1971). Female home range from a variety of geographic regions averages 5-60 km2 and males typically use areas larger areas from 15-220 km2 (Sunquist & Sunquist 2002). Home ranges in Namibia have been observed from 211-440 km2 (Marker & Dickman 2005). In Israel, home ranges for male caracals were 220 km2 +/- 132 km2, approximately four times larger than those of adult females (Weisbein & Mendelssohn 1990). A radio-collared male from Saudi

Arabia used a home range of 1116 km2 (Van Heezik & Seddon 1998). Ranges may expand or contract seasonally in some locales (Marker & Dickman 2005). Like most cats, caracal home ranges are a function of habitat type, habitat quality, resource availability and intraspecific competition (Sunquist & Sunquist 2002). Dispersal distances of young males are unknown.

Wide-ranging behavior of caracals enables them to recolonize areas following extensive removal efforts (Marker & Dickman 2005). Territories of both sexes are patrolled, defended, and demarcated with urine and feces (Avenant & Nel 1998). Male territories overlap at higher rates than females (Weisbein & Mendelssohn 1990; Marker & Dickman 2005), thus females may 33

be strongly defending territories from conspecifics. Male caracals move at a higher rate, further distances and have larger home ranges compared to females (Moolman 1986; Avenant & Nel

1998). Male caracals have been recorded wandering 5-15km daily, whereas females moved 2.5-

10km daily (Weisbein & Mendelssohn 1990).

Reproduction-

Caracals reproduce year-round, but reproduction is reduced in winter months (Bernard &

Stuart 1987). Reproductive peaks may be driven by female condition and nutritional status.

Juveniles have been observed from November-July in Zambia (Smithers 1983). Caracals that prey on livestock have been hypothesized to more often in good condition to reproduce (Bernard

& Stuart 1987), but no evidence supports this claim (A. Neils, personal observation). Female caracals may mate with multiple males during estrus (Weisbein & Mendelssohn 1990).

According to Botswana bushmen, paired caracals remain together for four days while mating

(Smithers 1971). Females use burrows made by other animals for denning, such as aardvarks

(Orycteropus afer) and African crested porcupines (Hystrix cristata) (Smithers 1983; Stuart &

Stuart 1985). Gestation is 69-78 days (Stuart & Stuart 1985; Bernard & Stuart 1987; Weisbein

& Mendelssohn 1990), and litters range from 2-5 kittens (Smithers 1983; Weisbein &

Mendelssohn 1990; A. Neils, personal observation). Young are born blind and deaf, with eyes opening at 0.5-1.5 weeks (Stuart & Stuart 1985). Incisors are visible at 10 days, full dentition visible at round 50 days and dentition is complete at 10 months (Stuart & Stuart 1985). At 2.5 weeks kittens begin vocalizing and by 4 weeks claws are fully retractable. From 3.5-8 weeks kittens begin exploring outside of the den and begin to consume their first solid foods. By 10

weeks kittens are active, mobile and able to defend themselves (Stuart & Stuart 1985). Weaning occurs from 15-20 weeks (Stuart & Stuart 1985). Kittens are lighter or grayer in color and more spotted than adults. Male caracals reach sexual maturity from between 7-12 months and females reach sexual maturity at 9-14 months (Bernard & Stuart 1987). Breeding does not likely occur for several more months unless there is an absence of competition in the population. Longevity records are exceedingly scarce, coming only from captive animals, but may be around 16 years

(Smithers 1983). This number may be markedly smaller for wild individuals (personal observation).

Conservation Status-

Caracals are considered rare, threatened, endangered or extirpated throughout most of their range (Nowell & Jackson 1996; Sunquist & Sunquist 2002). Two principal threats to caracals range-wide are habitat destruction and human persecution (Sunquist & Sunquist 2002).

Asian caracal populations are included in CITES Appendix I, and African populations are included on Appendix II (www.iucnredlist.org). In Kenya, Tanzania, Mozambique, Uganda,

Sudan and Niger the principal threat to caracals is increased hunting and poaching brought about by political instability (Sunquist & Sunquist 2002). Anthropogenic development including forest clearing, mining and hydro-electric projects reduce available habitat and encourage illegal poaching. Though caracal pelts generally hold very low commercial value, spotted belly fur of caracals is becoming increasingly valuable in Asian fur markets (Sunquist & Sunquist 2002).

Restrictions in big cat hunting in southern African has amplified the demand for caracal trophy hunting in Namibia (personal observation). Hunting caracals is legally prohibited in

Afghanistan, Algeria, Egypt, India, Iran, Israel, Jordan, Kazakhstan, Lebanon, Morocco,

Pakistan, Syria, Tajikistan, Tunisia, Turkey, Turkmenistan, and Uzbekistan (Nowell &Jackson

1996). In sub-Saharan Africa, caracals are protected in about half their range (Nowell & Jackson

1996). In Namibia and South Africa however, caracals are persecuted as a threat to livestock and legally classified as ‘vermin’, which permits landowners to kill without restriction.

The relationship between humans and caracals is best defined in terms of human-wildlife conflict. In southern Africa, all carnivores were systemically destroyed because of predation on domestic stock when the first European settlers arrived around 1820 (Pringle & Pringle 1979).

Following the removal of larger carnivores like leopards, lions, and hyaena, caracals became the apex predator in many regions and the focus of eradication campaigns. Caracals are considered the greatest predatory threat to small stock such as sheep and goats in southern Africa (Stuart &

Hickman 1991; Mukherjee et al. 2004; Berger 2006) and occasional surplus killing of livestock is reported (Skinner 1979; Stuart 1986; Braczkowski et al. 2012). Thus, caracals are killed rampantly in southern Namibian as perceived depredators of livestock.

Insufficient data exist on population trends for sub-Saharan caracal populations. Prior to the 1990s, most research programs on caracals consisted of finding effective means of exterminating the species (Stuart 1986). From 1931-1952, over 2,200 caracals per year were killed in control operations in the Karoo, South Africa (Stuart 1982). Farmers in Namibia reported killing up to 2,800 Caracals in a single year (Nowell & Jackson 1996). This pace of persecution may have lasted for decades, as many regions in southern Africa are saturated with livestock farms whose managers persistently destroy caracals. Caracals were once thought to have the capacity to withstand indefinite persecution pressure (Stuart 1981; Stuart 1986), but 36

eventual long-range declines are forecasted (Visser 1976; A. Neils, personal observation).

Caracal persecution on southern Namibian farmlands caused significant changes to caracal behavior and population dynamics and is not sustainable (A. Neils, personal observation). In some areas, caracal persecution has been implicated as the cause for an increase in prey numbers, particularly hyrax, which can lead to significant loss of forage and therefore financial loss to farmers (Palmer & Fairall 1988; Melville et al. 2004).

Unfortunately, certain economic industries are often in direct competition such as livestock production and conservation/ ecotourism. Although predator control efforts utilized by livestock producers have been successful in terms of the number of carnivores removed, they often fail to prevent declines in the livestock industry (Berger 2006). In Namibia, farmers pressure each other to kill predators, even if the threat is merely perceived (A. Neils, personal observation). Caracal persecution is difficult to curtail when the goal of most farmers is to ultimately realize eradication. Under the current legislation of Namibia, landowners act within their legal rights to achieve this end.

Caracals are at a crossroads and their future is uncertain, even in the stronghold of southern Namibia. Caracals require immediate scientific surveys and population censuses in all range countries. In Namibia, long-term viability of caracals should incorporate expanses outside of protected parks, as populations of smaller felids are low in most protected reserves with higher densities of larger heterospecifics (Caro 1987; Stewart & Wilson 1988). Caracals should be removed as ‘vermin’ in South Africa and Namibia, since these countries represent the last stronghold for the species and there is no scientific evidence to warrant this listing. If caracals are reclassified from ‘vermin’ status, they may still have the capacity to rebound if persecution 37

pressure is relaxed. Better scientific understanding of the relationship between human activity and caracal population dynamics is required to prevent long range declines.

Literature Cited

ACKERMAN, B.B., F.G. LINDZEY, AND T.P. HEMKER. 1984. Cougar food habits in southern Utah. The Journal of Wildlife Management: 147-155.

AVENANT, N.L, AND J.A.J. NEL. 1998. Home-range use, activity, and density of caracal in relation to prey density. African Journal of Ecology 36:347-359.

BERGER, K.M. 2006. Carnivore-Livestock Conflicts: Effects of Subsidized Predator Control and Economic Correlates on the Sheep Industry. Conservation Biology 20:751–761.

BERNARD, R.T.F., AND C.T STUART. 1987. Reproduction of the caracal Felis caracal from the Cape Province of South Africa. South African Journal of Zoology 22:177-182.

BISWAS, S., AND K. SANKAR. 2002. Prey abundance and food habit of tigers (Panthera tigris tigris) in Pench National Park, Madhya Pradesh, Indian Journal of Zoology (Lond.) 256:411–420.

BOTHMA, J.DU P. 1965. Random observations of the food habits of certain Carnivora of South Africa. Fauna and Flora 16:16-22.

BRACZKOWSKI, A., L. WATSON, D. COULSON, J. LUCAS, B. PIESER, AND M. ROSSI. 2012. The diet of the caracal, Caracal caracal, in two areas of the southern Cape, South Africa as determined by scat analysis. South African Journal of Wildlife Research 42:111-116.

BRAND, D.J. 1989. The control of caracal (Felis caracal) and baboons (Papio ursinus) in the Cape Province with the help of mechanical means. Unpublished M. Sc. thesis. University of Stellenbosch, Stellenbosch.

CARO, T.M., 1987. Cheetah’s mothers’ vigilance looking out for prey or for predators. Behavioral Ecology Sociobiology 20:351-362.

DE SMET, K.J.M. 1989. Studie van de verspreiding en biotoopkeuze van de grote mammalia in Algerije in het kader van het natuurebehoud. Ph.D. dissertation, Rijks University, Gent, Holland.

DU PLESSIS, J.J., N.L., AVENANT, AND H.O. DE WAAL. 2015. Quality and quantity of scientific information available on black-backed jackals and caracals: contributing to human-predator conflict or management? African Journal of Wildlife Research 45:138- 157.

GAILLARD, N. 1969. Sur la presence du chat dore (Felis aurata Temminck) et du caracal (Felis caracal Schreber) dans le sud du Senegal. Mammalia 33:350-351.

GROBLER, J.H. 1981. Feeding behavior of the caracal Felis caracal Schreber 1776 in the Mountain Zebra National Park. South African Journal of Zoology. 16:259-262.

HAPPOLD, D.C.D. 1987. The mammals of Nigeria. Oxford: Clarendon Press.

HEPTNER, V.G., AND A.A. SLUDSKII. 1992. Mammals of the Soviet Union. Vol 2.2, Carnivora (Hyeanas and cats). R.S. Hoffmann. Washington, DC.

HUFNAGL, E., 1972. Libyan mammals. Stoughton, WI: Oleander Press.

JOHNSON, W.E., E. EIZIRIK, P. PECON-SLATTERY, W.J. MURPHY, A. ANTUNES, E. TEELING, AND S.J. O’BRIEN. 2006. The Late Miocene Radiation of Modern Felidae: A Genetic Assessment. Science 311:73-77.

KOHN, T.A., AND T. NOAKES. 2013. Lion (Panthera leo) and caracal (Caracal caracal) type IIx single muscle fibre force and power exceed that of trained humans. Journal of Experimental Biology 216:960-969.

KOK, O.B., AND J.A.J. NEL. 2004. Convergence and divergence in prey of sympatric canids and felids: opportunism or phylogenetic constraint? Biological Journal of the Linnean Society 83:527-538

KOWALSKI, K., AND B. RZEBIK-KOWALSKI. 1991. Mammals of Algeria. Warsaw: Polish Academy of Sciences.

KINGDON, J. 1977. East African mammals. Vol. 3A (Carnivores). Chicago, University of Chicago Press.

LLOYD, P.H., AND J.C.G. MILLAR. 1981. A questionnaire survey of some of the larger mammals of the Cape Province. Bontebok 1:1-58.

LYNCH, C.D., 1983. The mammals of the Orange Free State. Memoirs van die Nasionale Museum Bloemfontein 18:218.

MALLON, D., AND K. BUDD. 2011. Regional Red List status of carnivores in the Arabian Peninsula. IUCN, Cambridge and Gland and Environment and Protected Areas Authority, Sharjah.

MARKER, L., AND M. DICKMAN. 2005. Notes on the spatial ecology of caracals (Felis caracal) with particular reference to Namibian farmlands. African Journal of Ecology 43, 73-76.

MELVILLE, H.I.A.S, J.DU P. BOTHMA, AND M.G.L. MILLS. 2004. Prey selection by caracal in Kgalagadi Transfrontier Park. South African Journal of Wildlife Research 34:67-75.

MELVILLE, H.I.A.S., AND J.DU P. BOTHMA. 2006. Using spoor counts to analyse the effect of small stock farming in Namibia on caracal density in the neighboring Kgalagadi Transfrontier Park. Journal of arid Environments 64:436-447.

MENDELSSOHN, H. 1989. Felids in Israel. Cat News 10:2-4.

MOOLMAN, L.C. 1984. ‘n vergelyking van die voedingsgewoontes van die rooikat Felis caracal binne en buite die Bergkwagga Nasionale Park. Koedoe 27:121-129.

MOOLMAN, L.C. 1986. Aspects of the ecology and behavior of the caracal Felis caracal Schreber 1776 in the Mountain Zebra National Park and on the surrounding farms. Master’s thesis, University of Pretoria, Pretoria.

MUKHERJEE, S., S.P. GOYAL, A.J.T. JOHNSINGH, AND M.R.P. LEITE PITMAN. 2004. The importance of rodents in the diet of jungle cat (Felis chaus), caracal (Caracal caracal), and golden jackal (Canis aureus) in Sariska Tiger Reserve, Rajasthan, India. Journal of Zoology (Lond) 262:405-411.

NADER, I.A. 1984. A second record of the caracal lynx Caracal caracal schmitzi (Matschie, 1912) for Saudi Arabia (Mammalia: Carnivora). Mammalia 48:148-150.

NOWELL, K., AND P. JACKSON. 1996. Wild cats: status survey and conservation action plan. Gland: IUCN 382.

PALMER, R., AND N. FAIRALL. 1988. Caracal and African wildcat diet in the Karoo National Park and the implications thereof for hyrax. South African Journal of Wildlife Research. 18:30-34.

PIENAAR, U.DE V. 1969. Predator-prey relationships amongst the large mammals of the Krugger National Park. Koedoe 12:108- 183.

POCOCK, R.I. 1939. The fauna of British India, including Ceylon and Burma. Mammalia: Primates and Carnivora. New Delhi.

PRINGLE, J.A., AND V.L. PRINGLE. 1979. Observations on the lynx Felis caracal in the Bedford district. South Africa Journal of Zoology 14:1-4. 41

PRATER, S.H. 1971. The book of Indian animals, 3rd ed. Bombay: Bombay Natural History Society.

REDDY, H.S., C. SRINIVASULU, AND K.T. RAO. 2004. Prey selection by the Indian tiger (Panthera tigris tigris) in Nagarjunasagar Srisailam Tiger Reserve, India. Mammalian Biology 69:384–391.

ROBERTS, T.J. 1977. The Mammals of Pakistan. London: Ernest Benn.

SICURU, F.L. 2011. Evolutionary trends on extant cat skull morphology (Carnivora: Felidae): a three-dimensional geometrical approach. Biological Journal of the Linnean Society 103:176-190.

SINGH, R., Q. QURESHI, K. SANKAR, P. KRAUSMAN, AND S.P. GOYAL. 2014. Population and habitat characteristics of caracal in semi-arid landscape. Indian Journal of Arid Environments 103:92-95.

SKINNER, J.D. 1979. Feeding behavior in caracal Felis caracal. Journal of Zoology (London) 189:523-525.

SMITHERS, R.H.N. 1971. The mammals of Botswana. Museum Memoir no. 4. Salisbury: The Trustees of the national Museums of Rhodesia.

SMITHERS, R.H.N. 1983. The Mammals of the Southern African Subregion. Pretoria: University of Pretoria.

SMITHERS, R.H.N, AND V.J. WILSON. 1979. Checklist and atlas of the mammals of Zimbabwe Rhodesia. Memoir No. 9, National Museums and Monuments, Salisbury.

STUART, C.T. 1981. Notes on the mammalian carnivores of Cape Province, South Africa. Bontebok 1:1-58.

STUART, C.T. 1984. The distribution and status of Felis caracal Schreber, 1776. Saugetierkundliche Mitteilungen 31:197- 203.

STUART, C.T. 1986. The Incidence of Surplus Killing by Panthera pardus and Felis caracal in Cape Province, South Africa. Mammalia 50:556-558.

STUART, C.T., AND G.C. HICKMAN. 1991. Prey of caracal, Felis caracal, in two areas of Cape Province, South Africa. Bontebok 1: 1-58. .

STUART, C.T., AND T.D. STUART. 1985. Age determination and development of foetal and juvenile Felis caracal Schreber 1776. Saugetierkundliche Mitteilungen 32, 217-229.

STUART, C. AND T. STUART. 2013. Caracal Caracal caracal: Mammals of Africa 5:174-179.

STUART, C.T., AND V.J. WILSON. 1988. The cats of Southern Africa. Zimbabwe: Chipangali Wildlife Trust.

SUNQUIST, M., AND F. SUNQUIST. 2002. Wild cats of the world. Chicago: University of Chicago Press.

THORN, M., M. GREEN, P. BATEMAN, E.Z. CAMERON, R. YARNELL, AND D. SCOTT. 2010. Comparative performance of sign surveys, spotlighting, and audio playbacks in a landscape-scale carnivore survey. South African Journal of Wildlife Research 40:77-86.

VAN HEEZIK, Y.M., AND P.J. SEDDON. 1998. Range size and habitat use of an adult male caracal in northern Saudi Arabia. Journal of Arid Environments 40:109-112.

VILJOEN, S., AND D.H.S. DAVIS. 1973. Notes on the stomach contents analyses of various carnivores in southern Africa (Mammalia: Carnivora). Annals of the Transvaal Museum 28:353-362.

VISSER, J. 1976. Status and conservation of the smaller cats of southern Africa. In: The World’s Cats, vol. 3 ed. R.L. Eaton. Seattle: Carnivore Research Institute:60-66.

WEISBEIN, Y., AND H. MENDELSSOHN. 1990. The biology and ecology of the caracal Felis caracal in the northern Aravah Valley of Israel. Cat News 12:20-22.

Figures

Figure 1. Map of current geographic range of Caracal caracal based on available data (2018).

APPENDIX A: DIET OF CARACALS (CARACAL CARACAL) ON NAMIBIAN FARMLANDS

Aletris M. Neils

(In the format of Journal of Mammalogy)

Aletris M. Neils*, [email protected]

Diet of Caracals (Caracal caracal) on Namibian Farmlands

Running Head: Namibian Caracal Diet

Diets were determined for caracals on working farms in southern Namibia from 2010 to 2016. Four different ecosystems were surveyed including Karoo/Karasberg, Kalahari grassland, sparse shrubland, and Acacia thornbrush. Relative frequencies of caracal prey items were calculated from analysis of 202 caracal stomachs, 688 kill sites and 250 caracal scats (N=1140 samples). A total of 106 species were recorded in Namibian caracal diet. Mammalian prey made up 83.2% of diet and ranged from 62.1% -

89.7% across the four ecosystems. Artiodactyls, rodents, lagomorphs, birds and sympatric carnivores were important components of Namibian caracal diet. Domestic stock comprised only 2.1% of caracal diet despite their abundance throughout the study areas. Proportions of major mammalian and avian taxa varied within each ecosystem. Prey availability and prey vulnerability are likely influencing factors that determine relative frequencies of prey items in caracal diet across ecosystems, but also the presence of larger sympatric felids is suspected to play a role in determining the realized predatory niche of caracals.

Keywords: Diet, Caracal caracal, farmlands, Namibia, predation, scat, stomach contents, wildlife conflict

*Aletris M. Neils: Conservation CATalyst, 3755 West Driscol Lane, Tucson, AZ 85745

Conﬂicts between wildlife and people pose a significant challenge to conservationists

(Woodroffe & Ginsberg 2005). Human–wildlife conﬂict (HWC) arises when a species poses a direct and recurring threat to community livelihood. HWC occurs in many different habitats but rangelands have always been particularly prone to conﬂicts due to the proximity of carnivores, prey, people and livestock (Zimmerman 2010). Much of Namibia’s agricultural land (44%) is utilized for livestock production (Schneider 1994). Namibian farmlands support a combination of domestic livestock (cattle, goats, sheep, and poultry) and game but are also home to the majority of Namibia’s wildlife (Marker et al. 2003).

Carnivores on the landscape are a common target of blame for suboptimal stock production and unexpected stock losses and death (Kellert et al. 1996; Treves and Karanth 2003; Musiani and

Paquet 2004). In Namibia and most of southern Africa, caracals (Caracal caracal) are commonly regarded as vermin due to occasional predation on small stock (Avenant and Nel 1998). Namibian farmers considered caracals to be the top threat to small stock and the fourth greatest threat to their cattle (Stein 2006). Magnitude of HWC can be real or perceived, but often results in persecution efforts of predators (Zimmerman et al. 2010). Persecution, defined as the persistent killing, chasing or other harassment of a species with the goal of eradication, is common among commercial ranchers that lose livestock to predation (Zimmerman et al. 2010). Southern African caracals face tremendous retaliatory persecution. From 1931-1952, over 2,200 caracals per year were killed in control operations in the Karoo, South Africa (Stuart 1981). Similarly, farmers in

Namibia reported killing up to 2,800 caracals in 1981 alone (Nowell and Jackson 1996). This pace

of persecution may have lasted for decades on a regional scale (Stuart & Hickman 1991; Stuart &

Stuart 2013) although no recent data exist.

In order to understand the causation of rangeland HWC, accurate data on the diet consumed by carnivores is essential (Karanth et al. 2004; Treves and Karanth 2003). HWC often occurs where natural prey is depleted and livestock provides an alternative food source (Woodroffe &

Ginsberg 1998). Although in some areas caracals may take livestock (Skinner 1979; Stuart 1986), caracals predominantly prey upon small antelope, rodents, rock hyrax (Procavia capensis), and birds (Grobler 1981; Palmer and Fairall 1988). Stock predation rates appear correlated with low natural prey densities, drought, and colder weather (Palmer and Fairall 1988; Melville et al. 2004;

Melville and Bothma 2006). The threat that caracals pose to domestic livestock could be less than is commonly perceived.

A thorough understanding of caracal natural history and dietary habits are lacking yet required to examine the current conflict. Documenting prey composition is a common objective of research studies involving predators as dietary studies provide key data to understand predator ecology. Information on predation is among the most important aspects of large carnivore ecology from a conservation and management perspective because managers are often faced with decisions that influence both predator and prey (Karanth et al. 2004; Musiani and Paquet 2004; Rominger et al. 2004). The purpose of this study was to examine caracal diet on southern Namibian farmlands, to determine extent of livestock depredation, and whether predation varies by habitat. Different methods (stomach contents, kill sites, and scats) are compared to determine methodology- associated biases (Meinzer et al. 1975; Ackerman 1984; Helldin 2000; Andersone and Ozoliņs

2004; Zabala and Zuberogoitia 2003; Bacon et al. 2011). 48

METHODS

Study Area.- This research was conducted on privately owned farmlands throughout

Namibia. Telemetry research was concentrated in the southern half of Namibia, from the village of Dordabis (70km southwest of Windhoek) south to the Groot Karas Mountains (130km southwest of Keetmanshoop) (Fig. 1). The region is extremely arid with highly fluctuating and unpredictable rainfall (Quan et al. 2004). Livestock farming with sheep, goats and some cattle is the predominant land use. Large carnivores including lions (Panthera leo), leopards (Panthera pardus), spotted hyenas (Crotua crotua), and cheetah (Acinonyx jubatus) have been effectively extirpated from southern Namibia due to persecution from livestock producers, thus caracals now serve as the apex predator in the southern half of the country. In southern Africa caracals typically inhabit arid and semi-arid shrub savannahs, open grasslands, Acacia thornbrush, rocky scrublands and mixed riverine habitats (Smithers 1971). To address habitat differences in caracal prey availabilty, this research was conducted in four study areas representing distinct ecosystems

(Karoo/ Karasberg, Kalahari grassland, sparse shrublands and Acacia thornbrush) (Fig. 1). All four ecosystems are defined by variations in soil, topography, vegetation structure. Research priority for this study was given to areas with the highest caracal–farmer conflict.

Kalahari grassland- Kalahari grassland plains stretch from central Namaland in the north up to the Blydverwagt Plateau south of Ariamsvlei in the south (Fig. 1). Kalahari grassland is a savanna ecosystem characterized by the dominance of grasses. Kalahari grassland has a relatively high mean vegetation cover (58%) (Okitsu 2010). Though the general appearance of Kalahari grassland is similar to that of a typical grass-dominated savanna, variability in vegetation is much more localized. Kalahari grassland contains a relatively higher amount of woody plants, both 49

deciduous and evergreen (Okitsu 2010). Ground is dominated by annual and perennial grasses with a variety of scattered large Acacia spp. The general structure of the habitat varies between tall open shrubland to a low open shrubland, with occasional low open to sparse Acacia thornbrush included (Strohbach 2001). Topography is variable and based on scattered small, hilly dune systems (Strohbach 2001). Kalahari grassland soils are sandy and stone cover is relatively low.

Karoo/ Karasberg- Karoo/ Karasberg habitat is found in the vicinity of Helmaringhausen,

NA and is located on the edge of the Karas Mountains along the western edge of the greater Nama-

Karoo biome (Fig. 1). Nama-Karoo is a complex of extensive plains, dominated by dwarf shrubs

(generally <1 m tall) intermixed with grasses, succulents, geophytes and annual forbs. Small trees occur only along drainage lines or on rocky outcrops that provide the right microclimate. The

Karoo/ Karasberg ecosystem is structured by the granitic inselbergs of the Great Karas Mountains with abundant rock outcrops and boulders across the landscape. Vegetation is short open shrubland with tree cover at 0.32 % (±1.7 %), shrub cover at 0.99 % (±2.8 %), dwarf shrub cover at 3.80 % (±4.9 %), grass cover at 5.31 % (±4.8 %) and herb cover at 1.39 % (±1.7 %) (Strohbach

2001). Vegetative cover in the Karoo/ Karasberg ecosystem is highly variable and a function of microhabitat. This ecosystem exhibits relatively high amounts of endemism and rich succulent diversity. Dominant plant life includes grasses such as Themeda triandra, Digitaria eriantha,

Tragus koelerioides and various dwarf shrubs such as Pentzia and Pteronia spp. (Mucina et al.

2006).

Sparse shrublands- Sparse shrubland habitat is also found within the greater Nama-Karoo biome. This study area is located in the vicinity of Koes, NA and falls within an area known as the Koes panveld (Fig. 1). Vegetation in this ecosystem is generally a low open shrubland in some 50

places becoming a tall open shrubland. Tree cover is extremely low and all trees exist in scrub form. Rocky ground cover is ample in the form of limestone, and soil is primarily hard kalk.

Ground is predominantly bare; shrub cover is 1.16 % (±1.7 %); dwarf shrub cover is 2.11 % (±2.2

%), grass cover is 3.36 % (±4.6 %) and herb cover is 1.78 % (±2.3 %) (Strohbach 2001). The

Koes panveld is very flat with poorly defined drainage patterns, so many pans have formed in this area.

Acacia thornbrush.- The Acacia thornbrush ecosystem (Fig. 1) is the dominant vegetation type over the central region of Namibia. Vegetation is typical of xeromorphic thornbush savanna with numerous woody plant genera. The Acacia thornbrush ecosystem varies in different parts of the area, but the characteristic feature is grassland with trees and bigger shrubs in dense or open clumps of varying size. A diversity of Acacia spp. dominate large parts of this region, including

Acacia reficiens, A. hebeclada, A. mellifera, A. erubescens, A. fleckii, and in some areas A. tortilis heteracantha. Ziziphus mucronata is found throughout most of this zone as well. Dichrostachys,

Grewia, Terminallia, and Boscia are common in some localities as well as Lonchocarpus nelsii in the sandy substrates and Combretum apiculatum on limestone and rocky outcrops. Understory vegetation is relatively sparse, although ephemeral forbs are present following rains. This region has been extensively modiﬁed over the last century through anthropogenic causes compounded by climatic ﬂuctuation (Quan et al. 1994; Rhode 1997). Perennial grasses have been reduced throughout this area to such an extent that remnant patches of historic open savanna habitat exist only where livestock grazing has been limited (Meik et al. 2002).

Animal Trapping, Handling, and Immobilization.- Study animals used for this research included live-trapped and deceased caracals captured and donated by farmers. Killed caracals 51

included fresh, frozen, and desiccated specimens. An accurately represented cross-section of the population was assumed for this study due to the variety of methods used by southern Namibian farmers to kill caracals. These include shooting at night with spotlights (with and without predator- calling), steel gin traps/ leg-hold traps, cage traps, body snares, jackal canons (a wire-triggered gun typically rigged to fences and burrows), poisons, hunting dogs, and deliberate vehicular collisions.

Handling of animals followed the American Society of Mammalogist’s Animal Care and Use

Committee protocol (http://www.mammalsociety.org/committees/animal-care-and-use). The project was conducted under University of Arizona’s IACUC certification number 6809 and under research permit 1539 granted by the Namibian Ministry of the Environment.

After capture, caracals were immobilized with an injection of a 1:1 mixture of Tiletamine hydrochloride and Zolazepam hydrochloride (Telazol™). This was administered at 3.0- 3.5 mg/kg of estimated body weight via a hand injection using a jab stick. Blowpipe or remote injection gun was used to immobilize cats opportunistically caught in chicken coops, livestock pens, snares or leghold traps. VHF and GPS radio collars (Followit, Tellus GPS; Lindesburg, Sweden) were fitted around the neck of tagged cats. Collars weighed between 285-370 g (<3% caracal body mass) and were equipped with mortality sensors. Caracals recovered fully from anesthesia in transport cage and were released when alert, usually within two hours of sunset and within 500 m of capture location.

Monitoring.- Following release, radio-collared cats were monitored at least every three hours via GPS collars. At least once per week, cats were triangulated and approached on foot to within 1km, where store-on-board GPS points were downloaded using UHF. All locations where caracals spent >3 hours were forensically investigated. If a kill is too large to consume in one 52

sitting, caracals typically cache the remains under any available vegetation and return to feed until the carcass is finished (Grobler 1981). Thus, sites occupied for >6 hours could be accurately detected by radio-telemetry clusters as kill sites. Time spent at kill sites correlated positively with prey size. The success in locating a kill sites correlated positively with the length of time spent by caracals at the site and to prey mass. Small prey such as rodents, lagomorphs and birds were not as reliably located from point clusters as larger artiodactyls. However, drops of blood, rodent tails and feathers were often detected. Investigating locations where caracals spent >1 hour may have led to more kill sites documented from small prey, but this was much more cost and labor intensive.

Scats were often collected in the vicinity of kill sites.

Determining Diet.- To determine caracal diet, kill sites from collared caracals and stomach samples from deceased caracals were examined. Caracal scat was also collected from collared caracals as well as opportunistically. By utilizing three different methodologies to determine caracal diet, not only could a broader spectrum of ingesta be documented but a comparison of the three different techniques to the diet composition as a whole was possible.

Kill sites were primarily discovered by investigating clusters of radio collared caracal activity, although some kill sites were opportunistically examined from non-collared cats for further information on prey species in the diet. Opportunistically examining kill sites followed observations of caracal hunts, from following up on caracal kills reported by farmers, and/or by back-tracking caracal spoor that led to a kill site. Prey remains (including hair, skin, rumen or stomach, and bone fragments) were used to identify prey species. Large viscera are usually left untouched by caracals which aids in locating kill sites. Prey remains, including the location of bite marks and portions consumed, were used to determine whether the caracal had killed the animal 53

or was scavenging. For all prey location of rumen was identified as the kill site and was marked with handheld Garmin GPS units (accuracy 5-10 m). For smaller prey, the largest collection of any remains was marked as the kill site and marked with GPS. All available depredated livestock and game were documented and examined to determine cause of death. Prey remains were identified to species, age and sex when possible. Data were collected on kill specifics (location, habitat, stalking location, struggle, kill bite, drag distance), carcass mass consumed, and number of times the cat returned to feed.

Caracal scat was collected at kill sites and opportunistically on roads and trails while radio- tracking collared caracals. Caracal scats were distinguished from those of African wild cat by the larger size of caracal scat (> 18 mm) and random manner of deposit (Stuart and Hickman 1991).

Of the scats collected, at least 45 verified scats (i.e.- associated with another form of caracal sign such as tracks or kill) from each habitat type were randomly selected for diet analysis. Over 40 scats were required to reach an asymptote in analysis of sampling effectiveness (Braczkowski et al. 2012). Due to the higher number of sympatric felids (serval, cheetah, and leopard) in Acacia thornbrush habitat, only scats associated with verified caracal track, kill site, or those collected in a cage trap with a caracal were analyzed. A caracal prey reference collection was constructed from kill site remains. Bird and herpetofauna remains were identified with assistance from local ornithological and herpetological experts and using museum collections.

Statistical analysis.- Examining frequency of occurrence data to estimate prey composition for carnivores is a common method for describing predator diets (Schaller 1967; Sunquist 1981;

Rabinowitz 1989; Johnsingh 1992; Venkataraman et al. 1995, Reddy et al. 2004). Relative frequencies were calculated for each observed caracal prey species as well as grouped Orders of 54

the prey items represented in observed caracal diet. The percent contribution (relative frequency) is the total number of individual prey items per category as a percentage of total number of prey items, and this was tallied for all habitat types and all methods used (kill sites, scats and stomach contents). To examine differences in observed caracal diet as influenced by habitat type, a non- parametric chi-square test (using relative frequencies) was used to model the linear relationship between habitat type and prey type. To examine differences in southern Namibian caracal diet as a function of habitat, prey frequencies from each habitat type were compared to prey frequencies for all southern Namibian caracals.

Several authors report inconsistent representations of diet depending on methodology used for a variety of carnivore taxa (Meinzer et al. 1975; Ackerman 1984; Helldin 2000; Andersone and

Ozoliņs 2004; Zabala and Zuberogoitia 2003; Bacon et al. 2011). A non-parametric chi-squared test was also used to compare diet results per method per habitat type.

RESULTS

A total of 202 caracal stomachs, 688 kill sites and 250 caracal scats were analyzed (N=1140 samples). One hundred and six species were recorded in southern Namibian caracal diet

(Appendix A). Mammalian prey made up 83.2% of southern Namibian caracal diet (Table 1), ranging from 62.1% - 89.7% of caracal diet across the four habitat types (Table 2). Karoo/

Karasberg and Kalahari grassland showed the highest relative frequencies of mammalian prey

(89.7 and 89.2 respectively) (Table 2). Mammalian prey was dominated by Artiodactyla (39.4%), followed by Hyracoidea (12.5%), Rodentia (12.2%), Lagomorpha (8.9%), and Carnivora (5.9%)

(Table 1). Artiodactyla prey ranged from 9.4% in Acacia thornbrush habitat to 64.3% in Kalahari

grassland (Table 2), representing the biggest range per habitat for any taxa except for the

Hyracoidea, which were the dominant prey item in Karoo/ Karasberg (43.4%) but were only present in two habitats. Top prey species overall were steenbok (Raphicerus campestris), springbok (Antidorcas marsupialis), rock hyrax (Procavia capensis), and Lepus hares (2 spp.).

These prey items constituted over 68% of all southern Namibian caracal diet. Caracals consumed

12 different carnivore species, the most common being black-backed jackal (Canis mesomelas) and African wildcat (Felis silvestris lybica). Domestic stock (Ovis aries and Capra aegagrus hircus) represented 2.1% of southern Namibian caracal diet overall (0.9%-5.1% per habitat).

Avian species combined for 13.3% of caracal diet with Otidiformes (primarily northern black korhaan, Afrotis afraoides) and Galliformes (primarily helmeted guineafowl, Numida meleagris) having the highest representation (3.5% and 3.6% respectively). Across habitat types, avian prey ranged widely from 7.9% in Karoo/ Karasberg to 31.1% in Acacia thornbrush (Table 2). All herpetofauna (15 species) combined for 3.6% of caracal diet (Table 1), primarily represented by squamates, particularly rock monitors (Varanus albigularis) (Appendix A).

Methodology.- Distribution of prey was different among methods used (χ2 = 612, df = 62,

P < 0.00001). Presence of artiodactyl prey was best represented by kill sites; relative frequency of artiodactyls more than doubled compared to examining only stomach contents and scats (Table

3). The same trend held for carnivores and Hyracoidea but to a lesser extent (Table 3). Rodent prey was underrepresented using kill site analysis alone as rodent material is typically consumed in its entirety. Rodents were best documented using scat analysis (Table 3). Likewise, avian prey was best documented using stomach contents and scat as opposed to kill sites (Table 3), although this trend did not hold in the Acacia thornbrush habitat (Table 2). Lagomorphs were relatively 56

evenly represented with all methodologies, but, like rodents, lagomorphs were best documented using scat analysis (Table 3). Domestic stock had the highest frequency of occurrence using stomach content analysis (4.0%) (Table 3).

Karoo/ karasberg- Distribution of prey was significantly different among ecosystem types

(χ2 = 1156, df = 93, P < 0.00001). Caracal diet in Karoo/ karasberg was determined from 324 total samples (53 caracal stomachs, 218 kill sites, and 53 scats). Karoo prey frequencies were defined by a higher than average mammalian prey content (Table 4) but exhibited the second lowest (23.7%) artiodactyl prey for all habitats (Table 2). Most mammalian prey (43.4%) in the

Karoo consisted of rock hyrax, found in abundance in habitats with ample rock and boulder structure. Relative frequencies of rodent and lagomorph were the second lowest reported for all habitat types, and avian prey was the lowest at 7.9% overall. The only incidence of caracals taking

Chacma baboons (Papio ursinus) occurred in Karoo/ karasberg habitat, representing a novel caracal prey species not yet reported in the scientific literature.

Kalahari grasslands- Caracal diet in Kalahari grasslands was determined from 473 total samples (71 caracal stomachs, 337 kill sites, and 65 scats). Kalahari grassland prey frequencies were defined by the highest average mammalian prey content (89.2%) and the highest artiodactyl prey (64.3%) for all habitats (Table 2). By method, stomach contents revealed the highest relative frequency of livestock (74.8%) compared to stomachs and scats respectively (45.1%, 46.7%)

(Table 5). Carnivore predation was highest for all habitats (8.5%). Relative frequencies of rodent prey were tied with Karoo/ Karasberg for the lowest observed frequencies (7.6%), and lagomorph prey was the lowest reported for all habitat types (5.3%) (Table 2). Avian prey was the second lowest after Karoo/ Karasberg habitat (9.3%), and reptile prey was the lowest recorded for all 57

habitat types (1.5%) (Table 2). The only incidence of caracals taking aardvark (Orycteropus afer) occurred in Kalahari grassland habitat. Domestic stock predation (1.9%) was comparable to the overall average for all caracals (2.1%).

Sparse shrublands- Caracal diet in sparse shrublands was determined from 203 total samples (41 caracal stomachs, 114 kill sites, and 48 scats). Relative frequency for mammalian prey was 80.2%, slightly lower than pooled frequencies for all habitats (83.2%, Table 2).

Frequency of artiodactyl and avian prey in sparse shrublands was virtually the same as those reported for all southern Namibian caracals (Table 2). Rodent predation was slightly lower in sparse shrubland compared to pooled habitats (10.5% vs. 12.2%), while carnivore predation was slightly higher (7.2% vs. 5.9%). Relative frequencies for lagomorph prey in sparse shrublands were highest for all habitats (16.9%), as well as herpetofauna (6.8%) (Table 2). Lagomorph and herpetofauna prey in sparse shrublands were almost double those observed for all caracals.

Domestic stock predation was highest in sparse shrubland habitat, 5.1% compared to 2.1% for all caracals. By method, stomach contents had the highest relative frequency of livestock (8.1%) while scats revealed a lower relative frequency than the total (3.3% vs. 5.1%) (Table 6).

Acacia thornbrush- Caracal diet in Acacia thornbrush was determined from 140 total samples (37 caracal stomachs, 19 kill sites, and 84 scats). Caracal diet in Acacia thornbrush had the lowest content of mammalian prey at 62% (Table 2). Relatedly, avian ingesta were highest in

Acacia thornbrush compared to other habitats. Most mammalian prey consumed in Acacia thornbrush were small mammals (rodents and lagomorphs) while artiodactyls accounted for less than 10% (Table 2). Acacia thornbrush had the lowest relative frequencies of carnivores and the second highest frequency of herpetofauna after sparse shrublands (Table 2). Examining caracal 58

diet in Acacia thornbrush using kill sites alone underestimated the frequency of total mammalian prey (Table 7). This trend was due to inherently lower predation on ungulates and higher predation rates on small mammals. Lagomorphs were evenly represented in Acacia thornbrush by all methodologies (Table 7). Higher bird predation was observed in Acacia thornbrush (Table 7).

Caracals preyed more upon Galliformes avifauna in Acacia thornbrush than in other ecosystems, an Order represented primarily by helmeted guineafowl and francolin (Pternis spp.). These species represented approximately half of the avian material consumed by caracals in Acacia thornbrush

(Table 7). These birds are relatively large and would leave substantial feather material on the ground at kill sites following an attack and during processing. Caracal predation on Otidiformes avifauna (primarily represented by northern black korhaans, Afrotis afraoides) was not as high as in sparse shrublands, likely a function of availability and differences in species composition between these two habitats.

DISCUSSION

Felid prey preference is influenced by many factors including prey size, prey availability

(density and scarcity) and prey vulnerability (Schaller 1972; Carbone et al. 1999; Hayward and

Kerley 2005). Caracals are thought to be opportunistic predators (Funston et al. 2001; Radloff and

Du Toit 2004) and are documented predating a broad range of prey species. Examination of southern Namibian caracal stomach contents, kill sites and scats in this study revealed a total of

106 species including ungulates, hyraxes, lagomorphs, rodents, birds, carnivores and herpetofauna.

Namibian caracals exhibit generalist feeding behavior (Grobler 1981; Moolman 1986; Palmer and

Fairall 1988; Skinner 1979; Smithers 1983; Stuart and Hickman 1991; Weisbein and Mendelssohn

1990) and caracal habitat use is positively correlated with density of mammalian prey (Avenant and Nel 1998) and caracal foraging is best described as mammal-based searches, prey that caracals and other cat species are phylogenetically adapted to capture (Kok and Nel 2004). Caracal diet varies by region, habitat, or locale (Moolman 1986). Caracals in Kalahari grassland and sparse shrubland showed a stronger preference for artiodactyl prey, while mammalian prey in the Karoo consisted primarily of rock hyrax. Caracal in Acacia thornbrush consumed primarily rodents.

Caracals in a South African national park primarily chose rock hyrax and mountain reedbuck

(Redunca fulvorufula) with mammals contributing 93.8% of the total ingesta (Grobler 1981).

Caracal prey in Karoo National Park consisted of rodents (39%), artiodactyls (28%), hyrax (22%), lagomorphs (19%), arthropods (17%), and birds and carnivores (2% respectively) (Palmer and

Fairall 1988). Though caracals consume a high percentage of mammalian prey, the percent composition of mammalian taxa varies across habitats.

Competitive release- Caracals commonly take prey more than 2-3 times their own mass, a trend most often observed in larger felids (Grobler 1981; Sunquist and Sunquist 2002). Caracals are sufficiently equipped to take large ungulates, as opposed to preying almost exclusively on small mammals as seen in the closely related serval (Leptailurus serval). Earlier caracal diet research noted an emphasis on rodents and other small prey species (Bothma 1965; Pienaar 1969;

Viljoen and Davis 1973), yet caracals took larger prey in other regions (Pringle and Pringle 1979;

Smithers 1983). Relative importance of large prey species versus small mammals seems to vary by region. In some ecosystems larger carnivores are extirpated resulting in caracals becoming the dominant predators (Sunquist and Sunquist 2002; Stuart and Stuart 2013). Apex carnivores including lions, leopards, spotted hyenas, and cheetah have been effectively removed from the 60

southern half of Namibia within the past few decades (Stuart and Stuart 2013). This was the case in all ecosystems in this study except for Acacia thornbrush, where leopards and cheetahs still exist in high densities (Caro 1987). Coexisting with larger felids, caracals may exhibit a different realized niche focused on smaller prey as opposed to large ungulates. Without large predators, caracals may have the ability to expand their niche and behave more like a higher-tiered predator.

This phenomenon is noted for ocelots and pumas in areas were jaguars have been extirpated

(Moreno et al. 2006). In Kalahari grassland, where relative frequencies of artiodactyl predation by caracals were markedly high, relative frequencies of small lagomorphs and rodents in the diet were lowest. The inverse occurred in Acacia thornbrush habitat where caracals consumed lower frequencies of artiodactyl prey and higher frequencies of small mammals. Trends are confounding in rocky habitats that have high densities of hyrax, as in the Karoo/ karasberg ecosystem, since caracals prefer hunting in rocky areas versus adjacent flats if available (Grobler 1981). Of caracal predation events upon springbok, 130 occurred in Kalahari grassland and 69 occurred in all other ecosystems combined. Gemsbok (Oryx gazelle) and blesbok (Damaliscus dorcas phillipsi) predation occurred only in Kalahari grassland, as did over half of the greater kudu (Tragelaphus strepsiceros) records. Caracals consumed more carnivores in Kalahari grassland and it was the only habitat where aardvark was a documented prey item. This record represents a novel caracal prey species in the scientific literature, a relatively large prey item in the same habitat where artiodactyl prey was most common.

In Acacia thornbrush, rodents formed the bulk of caracal diet. The same trend has been observed in other locales that also support leopard, cheetah or both (Bothma 1965; Pienaar 1969;

Viljoen and Davis 1973; Melville et al. 2004). In Acacia thornbrush, frequent avian prey may be 61

due to higher number of birds available, or to easier search and handling times, i.e. selection of prey as a function of prey vulnerability (Schaller 1972; Carbone et al. 1999; Hayward and Kerley

2005). Alternatively, sympatric felid competition may explain caracal diet shift in Acacia thornbrush (Moreno et al. 2006). An inventory of prey availability per habitat is recommended to determine whether differences in caracal diet are due to prey abundance or from sympatric competition.

Intraguild predation- In many ecological systems one or more species may act as both predator and competitor with other species at the same or similar trophic level. Intraguild predation, is considered an extreme case of interference competition (Fedriani et al. 2000). Caracal consumption of other carnivores as observed in this study is well documented (Lloyd and Millar

1981; Stuart 1984; Palmer and Fairall 1988; Melville et al. 2004; Braczkowski et al. 2012). Black- backed jackal (Canis mesomelas) and African wildcat (Felis silvestris lybica) were the most commonly encountered carnivores preyed upon by caracals, but ten other species were recorded as well (Appendix A). Caracal often prey on black-backed jackals (Melville et al. 2004), especially pups (Stuart and Wilson 1988), and jackals are anecdotally said to increase in areas where caracals are heavily persecuted (Stuart 1981). Examination of scat and stomach contents may also underestimate carnivore prey killed by caracals. In many instances very little if any carnivore meat was consumed. Hair and bone were rarely ingested, so without examining kill sites many instances of caracals killing other carnivores would not be documented.

Trends in Methodology- Kill site analysis derived from radio collar data can be used to study carnivores that take a wide range of prey species and sizes (Anderson and Lindzey 2003;

Zimmermann et al. 2007; Merrill et al. 2010; Ruth et al. 2010). Like other large carnivores, caracal 62

kill sites predominantly contained remains of large prey. Small prey such as rodents, and to a lesser extent lagomorphs and birds, were documented less from kill sites and best documented using scat analysis. While examining kill sites underestimates small prey, examination of scat alone may underestimate percentage of ungulates. Caracals were observed to feed off ungulates for up to six days, and scat collected from initial feedings of large prey contained choice meat and therefore little or no identifiable hair and/ or bone remains. Genetic analysis of scat would help correct for this but can be cost prohibitive. Examination of kill sites alone may accurately reflect diet of large carnivores that exclusively take large prey, such as African lions or tigers (Panthera tigris), but would become inherently less reliable for smaller carnivores that consume high percentages of small prey (Biswas and Sankar 2002; Zabala and Zuberogoitia 2003; Reddy et al.

2004).

Livestock depredation.- Caracals are considered the greatest predatory threat to small stock such as sheep and goats in southern Africa (Stuart and Hickman 1991; Mukherjee et al. 2004;

Berger 2006) and occasional surplus killing of livestock is reported (Skinner 1979; Stuart 1986;

Braczkowski et al. 2012). Thus caracals are killed rampantly in southern Namibian as perceived depredators of livestock (Stuart and Hickman 1991; A. Neils, personal observation). Although natural prey of caracals in southern Namibia consisted primarily of ungulates, livestock were consistently a small component of caracal diet. Only 4 out of 34 radio-collared caracals predated livestock. Since livestock were abundant in all study areas, caracals selected against this potential food source. Felids and other large predators may select against livestock if wild ungulate prey is abundant (Ackerman 1984; Biswas and Sankar 2002; Reddy et al. 2004). Stock predation by caracals correlates with low natural prey densities, drought, and cold weather (Palmer and Fairall 63

1988; Melville et al. 2004; Melville and Bothma 2006). Severity of livestock depredation by caracals is dependent on availability of wild prey and stock husbandry techniques (Stuart and

Stuart 2013). In a coastal South African park where natural prey numbers were anthropogenically diminished, 15% of caracal diet was ungulates (both native antelopes and domestic stock)

(Braczkowski et al. 2012). Many studies of caracal diet have been conducted near livestock farms with diminished natural prey, and thus had conflicts with caracals that struggled to meet their energetic requirements (Du Plessis et al. 2015). Several thousand sheep and goats are killed annually by caracals in the Cape Province of South Africa (Stuart and Hickman 1991). Stomach contents of caracals obtained from animal control operations contained livestock prey material

19% of the time (Stuart and Hickman 1991). Caracal stomachs were empty 37% of the time indicating that caracals may have struggled to acquire food resources in a prey-depleted study area

(Stuart and Hickman 1991). Some caracals on farms never prey on stock (Moolman 1986; Palmer and Fairall 1988). Livestock may serve as a last resort for caracals unable to obtain wild prey due to inexperience, old age, injury, or management practices that deplete prey.

Domestic stock had the highest frequency of occurrence when stomach content analysis was used. Sparse shrubland stomach contents had the highest relative frequency of livestock whereas scats revealed a lower relative frequency than the total. This discrepancy is explained by different digestibility of prey types and different passage rates of various prey parts (Bowland and

Bowland 1991). Most depredated livestock were sheep, and ingested sheep wool tends to remain in the stomach for long periods of time. In general, scat analysis risks underestimation of highly digestible prey relative to prey that leaves large portions of indigestible remains (Putman 1984).

Livestock predation by caracals may be over-represented by examining stomach contents alone, 64

which is often the sole method used to quantify caracal depredation of livestock using carcasses obtained from cull operations (Stuart and Stuart 1991). All available diet methodologies should be used when quantifying depredation.

Conﬂicts between wildlife and people pose a significant challenge to conservationists

(Woodroffe & Ginsberg 2005). Ecological, economic and social consequences of predator interactions occur in places where predators and stock producers share the same land and resources

(Breck 2004; Bagchi and Mishra 2006). Livestock depredation and subsequent predator control are a growing concern worldwide (Loveridge 2010), however, blanket condemnation of carnivores due to depredations from few problem animals is ecologically indefensible (Stuart 1984). In order to develop effective solutions to livestock carnivore conflicts, it must be determined why predators are attracted to livestock and what aspects of their environment, biology or ecology predisposes livestock depredation (Breck 2004). This requires an understanding of diet, behavior, and community ecology in relation to prey interactions.

Conﬂict resolution requires solutions based on best available ecological data for the species involved (Ajzen and Fishbein 1980). Comprehensive understanding of predator ecology can change attitudes towards predators (Mech 1996; Treves and Karanth 2003). HWC management must include outreach that influences perceptions of risk (Riley and Decker 2000), as education improves predator tolerance (Homern et al. 2007). Accurate incorporation of caracal diet data into management decisions is imperative. Studies that describe caracals as problematic predators of livestock in some locales (Stuart and Hickman 1991; Braczkowski et al. 2012) often are generalized range-wide; results from diet studies restricted to singular locales may not accurately depict dietary patterns of caracals on a regional scale. Caracal persecution on southern Namibian 65

farmlands caused significant changes to life history statistics and population dynamics and is not sustainable (A. Neils, unpublished data). Better scientific understanding of the relationship between carnivores and farmers is ultimately required to improve long-term coexistence of both entities.

ACKNOWLEDGMENTS

First and foremost, the author would like to express gratitude to local farming hosts,

Kiripotib Guest Farm (Hans and Claudia von Hase), Timm and Inez Miller, and Tyse and Annetta

Swart. I also acknowledge the many livestock farmers that contributed specimens and information;

I could not have done it without your cooperation and trust. Field research was conducted with meticulous detail in part by the exceptional Elin Gromberg and Magnus Nystrand. I am very grateful for field assistance from several passionate interns especially Elan Fox, Lily Harrison, A.

Veals, C. Cavanaugh, A. Pena, and S. Thomann and many others who contributed to the success of the project through the years. Necropsy protocols were developed with direction and guidance from Dr. Axel Hartmann. I thank John Koprowski for his guidance and multiple reviewers for constructive comments. This research was conducted with support from the Namibian Ministry of Environment and Tourism (under permit number 1539). Funding was provided by Fulbright,

Cheetah Kids, Crosby Family, B. Kay, M. Sloan, and contributions from many other donors.

LITERATURE CITED

ACKERMAN, B.B., LINDZEY, F.G., AND HEMKER, T.P. 1984. Cougar food habits in southern Utah. The Journal of Wildlife Management: 147-155.

ANDERSON JR, C.R., AND LINDZEY, F.G. 2003. Estimating cougar predation rates from GPS location clusters. The Journal of Wildlife Management: 307-316.

ANDERSONE, Ž., AND J. OZOLIŅŠ. 2004. Food habits of wolves Canis lupus in Latvia. Acta Theriologica 49: 357-367.

AVENANT, N.L, AND J.A.J. NEL. 1998. Home-range use, activity, and density of caracal in relation to prey density. African Journal of Ecology 36:347-359.

BACON, M.M., G.M. BECIC, M.T. EPP, AND M.S. BOYCE. 2011. Do GPS clusters really work? Carnivore diet from scat analysis and GPS telemetry methods. Wildlife Society Bulletin 35:409-415.

BAGCHI, S., AND C. MISHRA. 2006. Living with large carnivores: predation on livestock by the snow leopard (Uncia uncia). Journal of Zoology 268:217-224.

BERGER, K.M. 2006. Carnivore-livestock conflicts: effects of subsidized predator control and economic correlates on the sheep industry. Conservation Biology 20:751–761.

BISWAS, S., AND K. SANKAR. 2002. Prey abundance and food habit of tigers (Panthera tigris tigris) in Pench National Park, Madhya Pradesh, Indian Journal of Zoology (Lond.) 256:411–420.

BOTHMA, J. DU P. 1965. Random observations of the food habits of certain Carnivora of South Africa. Fauna and Flora 16:16-22.

BOWLAND, J.M., AND A.E. ROWLAND. 1991. Differential passage rates of prey components through the gut of serval Felis serval and black-backed jackal Canis mesomelas. Koedoe, 34:37-39.

BRECK, S.W. 2004. Minimizing carnivore-livestock conflict: the importance and process of research in the search for coexistence. People and predators: from conflict to coexistence. Island Press, Washington, D.C. 67

CARBONE, C., G.M. MACE, S.C. ROBERTS, AND D.W. MACDONALD. 1999. Energetic constraints on the diet of terrestrial carnivores. Nature 402:286–288.

CARO, T.M. 1987. Cheetah’s mothers’ vigilance looking out for prey or for predators. Behavioral Ecology Sociobiology 20:351-362.

DU PLESSIS, J.J., N.L. AVENANT, AND H.O. DE WAAL. 2015. Quality and quantity of scientific information available on black-backed jackals and caracals: contributing to human-predator conflict or management? African Journal of Wildlife Research 45:138- 157.

FEDRIANI, J.M., T.K. FULLER, R.M. SAUVAJOT, AND E.C. YORK. 2000. Competition and intraguild predation among three sympatric carnivores. Oecologia, 125:258-270.

FUNSTON, P.J., M.G.L. MILLS, AND H.C. BIGGS. 2001. Factors affecting the hunting success of male and female lions in the Kruger National Park. Journal of Zoology 253:419-431.

GIESS, W. 1971. A preliminary vegetation map of South West Africa. SWA Wissenschaftliche Gesellschaft.

GROBLER, J.H. 1981. Feeding behavior of the caracal Felis caracal Schreber 1776 in the Mountain Zebra National Park. South African Journal of Zoology. 16:259-262.

HAYWARD, M.W. AND G.I. KERLEY. 2005. Prey preferences of the lion (Panthera leo). Journal of Zoology 267:309-322.

HELLDIN, J.O. 2000. Seasonal diet of pine marten Martes martes in southern boreal Sweden. Acta Theriologica 45:409-420.

JOHNSINGH, A.J.T. 1992. Prey selection in three sympatric carnivores in Bandipur. Mammalia 56:517–526.

KARANTH, K.U., J.D. NICHOLS, N.S. KUMAR, W.A. LINK AND J.E. HINES. 2004. Tigers and their prey: predicting carnivore densities from prey abundance. Proceedings of the National Academy of Sciences 101:4854–4858.

KELLERT, S.R., M. BLACK, C.R. RUSH, AND A.J. BATH. 1996. Human culture and large carnivore conservation in North America. Conservation Biology 10:977-990.

KOK, O.B. AND J.A.J. NEL. 2004. Convergence and divergence in prey of sympatric canids and felids: opportunism or phylogenetic constraint? Biological Journal of the Linnean Society 83:527-538. 68

LOVERIDGE, A.J., S.W. WANG, L.G. FRANK AND J. SEIDENSTICKER. 2010. People and wild felids: conservation of cats and management of conflicts. Biology and conservation of wild felids 161.

LLOYD, P.H. AND J.C.G. MILLAR. 1981. A questionnaire survey of some of the larger mammals of the Cape Province. Bontebok 1:1-58.

MARKER, L.L., M.G.L. MILLS AND D.W. MACDONALD. 2003. Factors influencing perceptions of conflict and tolerance toward cheetahs on Namibian farmlands. Conservation Biology 17:1290-1298.

MECH, L.D. 1996. A new era for carnivore conservation. Wildlife Society Bulletin 24:397-401.

MEINZER, W.P., D.N. UECKERT, AND J.T. FLINDERS. 1975. Food niche of coyotes in the rolling plains of Texas. Journal of Range Management 28:22-26.

MEIK, J.M., R.M. JEO, J.R. MENDELSON III, AND K.E. JENKS. 2002. Effects of bush encroachment on an assemblage of diurnal lizard species in central Namibia. Biological Conservation, 106:29-36.

MELVILLE, H.I.A.S, J.DU P. BOTHMA, AND M.G.L. MILLS. 2004. Prey selection by caracal in Kgalagadi Transfrontier Park. South African Journal of Wildlife Research 34:67-75.

MERRILL, E., ET AL. 2010. Building a mechanistic understanding of predation with GPS-based movement data. Philosophical Transactions of the Royal Society of London B: Biological Sciences 365:2279-2288.

MORENO, R.S., R.W. KAYS, AND R. SAMUDIO JR. 2006. Competitive release in diets of ocelot (Leopardus pardalis) and puma (Puma concolor) after jaguar (Panthera onca) decline. Journal of Mammalogy 87:808-816.

MUCINA, L., ET AL. 2006. Nama-karoo biome. The vegetation of South Africa, Lesotho and Swaziland. Strelitzia, 19:324-347.

MUKHERJEE, S., S.P. GOYAL, A.J.T. JOHNSINGH, AND M.R.P LEITE PITMAN. 2004. The importance of rodents in the diet of jungle cat (Felis chaus), caracal (Caracal caracal), and golden jackal (Canis aureus) in Sariska Tiger Reserve, Rajasthan, India. Journal of Zoology (Lond) 262:405-411.

MUSIANI, M. AND P.C. PAQUET. 2004. The practices of wolf persecution, protection, and restoration in Canada and the United States. BioScience 54:50-60.

NORTON, P.M. AND A.B. LAWSON. 1985. Radio tracking of leopards and caracals in the Stellenbosch area, Cape Province. South African Journal of Wildlife Research 15:17-24.

OKITSU, S. 2010. Vegetation structure of the biomes in southwestern Africa and their precipitation patterns. African Study Monographs (Supplement) 40:77-89.

PALMER, R. AND N. FAIRALL. 1988. Caracal and African wildcat diet in the Karoo National Park and the implications thereof for hyrax. South African Journal of Wildlife Research. 18:30-34.

PIENAAR, U.DE V. 1969. Predator-prey relationships amongst the large mammals of the Krugger National Park. Koedoe 12:108- 183.

PRINGLE, J.A., AND V.L. PRINGLE. 1979. Observations on the lynx Felis caracal in the Bedford district. South Africa Journal of Zoology 14:1-4.

PUTMAN, R.J. 1984. Facts from faeces. Mammal Review 14:79-97.

QUAN, J., D. BARTON, AND C. CONROY. 1994. A Preliminary Assessment of the Economic Impact of Desertiﬁcation in Namibia. Research Discussion Paper 3:1–148. Directorate of Environmental Affairs, Windhoek.

RABINOWITZ, A. 1989. The density and behavior of large cats in a dry tropical forest mosaic in Huai Kha Khaeng Wildlife Sanctuary, Thailand. Natural History Bulletin of the Siam Society 37:235–251.

RADLOFF, F.G. AND J.T. DU TOIT. 2004. Large predators and their prey in a southern African savanna: a predator's size determines its prey size range. Journal of Animal Ecology 73:410-423.

REDDY, H.S., C. SRINIVASULU, AND K.T. RAO. 2004. Prey selection by the Indian tiger (Panthera tigris tigris) in Nagarjunasagar Srisailam Tiger Reserve, India. Mammalian Biology 69:384–391.

RHODE, R.F. 1997. Looking into the past: interpretations of vegetation change in western Namibia based on matched photography. Dinteria 25:121–149.

ROMINGER, E.M., H.A. WHITLAW, D.L. WEYBRIGHT, W.C. DUNN, AND W.B. BALLARD. 2004. The influence of mountain lion predation on bighorn sheep translocations. Journal of Wildlife Management 68:993-999.

RUTH, T.K., P.C. BUOTTE, AND H.B. QUIGLEY. 2010. Comparing ground telemetry and global positioning system methods to determine cougar kill rates. Journal of Wildlife Management 74:1122-1133.

SCHALLER, G.B. 1972. The Serengeti lion. University of Chicago Press, Chicago.

SCHNEIDER, H.P. 1994. Animal health and veterinary medicine in Namibia. Agrivet, Windhoek, Namibia.

SKINNER, J.D. 1979. Feeding behavior in caracal Felis caracal. Journal of Zoology (London) 189:523-525.

SMITHERS, R.H.N. 1971. The mammals of Botswana. Museum Memoir no. 4. Salisbury: The Trustees of the National Museums of Rhodesia.

SMITHERS, R.H.N. 1983. The Mammals of the Southern African Subregion. Pretoria: University of Pretoria.

STEIN, A. 2006. Interactions: Leopards vs. Brown Hyenas. Otjiwarongo, Namibia: Waterberg Carnivore Project Namibia.

STROHBACH, B.J. 2001. Vegetation Survey of Namibia; Namibia Wissensschaftliche Gesellschaft / Namibia Scientific Society Journal 49:93-124.

STUART, C.T. 1981. Notes on the mammalian carnivores of Cape Province, South Africa. Bontebok 1:1-58.

STUART, C.T. 1984. The distribution and status of Felis caracal Schreber, 1776. Saugetierkundliche Mitteilungen 1:197- 203.

STUART, C.T. 1986. The Incidence of Surplus Killing by Panthera pardus and Felis caracal in Cape Province, South Africa. Mammalia 50:556-558.

STUART, C.T., AND G.C. HICKMAN. 1991. Prey of caracal, Felis caracal, in two areas of Cape Province, South Africa. Bontebok 1: 1-58.

STUART, C., AND T. STUART, T. 2013. Caracal Caracal caracal: Mammals of Africa 5:174- 179.

STUART, C.T., AND V.J. WILSON. 1988. The cats of Southern Africa. Zimbabwe: Chipangali Wildlife Trust.

SUNQUIST, M.E. 1981. The social organisation of tigers (Panthera tigris) in Royal Chitwan National Park. Smithsonian Contributions Zoology 336:1–98.

SUNQUIST, M., AND F. SUNQUIST. 2002. Wild cats of the world. Chicago: University of Chicago Press.

TREVES, A., AND K.U. KARANTH. 2003. Human-carnivore conflict and perspectives on carnivore management worldwide. Conservation Biology 17:1491-1499.

VENKATARAMAN, A.B., R. ARUMUGAM, AND R. SUKUMAR. 1995. The foraging ecology of dhole (Cuon alpinus) in Mudumalai Sanctuary, southern India. Journal of Zoology (Lond.) 237:543–561.

VILJOEN, S. AND D.H.S. DAVIS. 1973. Notes on the stomach contents analyses of various carnivores in southern Africa (Mammalia: Carnivora). Annals of the Transvaal Museum 28:353-362.

WEISBEIN, Y., AND H. MENDELSSOHN. 1990. The biology and ecology of the caracal Felis caracal in the northern Aravah Valley of Israel. Cat News 12:20-22.

WOODROFFE, R. AND J.R. GINSBERG. 1998. Edge effects and the extinction of populations inside protected areas. Science 28:2126-2128.

ZABALA, J. AND I. ZUBEROGOITIA. 2003. Badger, Meles meles (Mustelidae, Carnivora), diet assessed through scat-analysis: a comparison and critique of different methods. Folia Zoologica 52:23-30.

ZIMMERMANN, B., P. WABAKKEN, H. SAND, H.C. PEDERSEN AND O. LIBERG. 2007. Wolf movement patterns: a key to estimation of kill rate? Journal of Wildlife Management 71:1177-1182.

ZIMMERMANN, A., N. BAKER, C. INSKIP, J.D. LINNELL, S. MARCHINI, J. ODDEN, G. RASMUSSEN AND A. TREVES. 2010. Contemporary views of human–carnivore conflicts on wild rangelands. In Wild rangelands: Conserving wildlife while maintaining livestock in semi-arid ecosystems: 129-151.

APPENDIX A

Comprehensive prey species list of southern Namibian caracals (Caracal caracal) as determined by kill site, stomach content and scat analyses from 2010-2016.

Mammalia Total N = 1140

Artiodactyla

Steenbok (Raphicerus campestris) 281

Springbok (Antidorcas marsupialis) 199

Klipspringer (Oreotragus oreotragus) 33

Common duiker (Sylvicapra grimmia) 10

Damara dik-dik (Madoqua kirkii) 6

Gemsbok (Oryx gazella) 2

Greater kudu (Tragelaphus strepsiceros) 7

Blesbok (Damaliscus dorcas phillipsi) 3

Warthog (Phacochoerus aethiopicus) 3

Domestic

Sheep (Ovis aries) 24

Goat (Capra aegagrus hircus) 5

Lagomorpha

Scrub hare (Lepus saxatilis) 61

Cape hare (Lepus capensis) 60

Red rock rabbit (Pronolagus spp.) 5

Carnivora

Aardwolf (Proteles cristata) 8

African wildcat (Felis silvestris lybica) 13

Small-spotted cat (Felis nigripes) 2

Small-spotted genet (Genetta genetta) 8

Black-backed jackal (Canis mesomelas) 18

Cape fox (Vulpes chama) 2

Bat-eared fox (Otocyon megalotis) 7

Banded mongoose (Mungos mungo) 1

Yellow mongoose (Cynictis penicillata) 5

Slender mongoose (Galerella sanguinea) 9

Zorilla (Ictonyx striatus) 6

Domestic

Domestic cat (Felis catus) 2 73

Rodentia

Springhare (Pedetes capensis) 37

Dassie rat (Petromus typicus) 13

Ground squirrel (Xerus spp.) 17

Whistling rat (Parotomys spp.) 6

Striped mouse (Rhabdomys pumilio) 4

Field mouse (Mus spp.) 4

Multimammate mouse (Mastomys spp) 12

Tree mouse (Thallomys spp.) 9

Rock rat (Aethomys spp.) 14

Short-tailed gerbil (Desmodillus auricularis) 9

Hairy-footed gerbil (Gerbillurus spp.) 11

Bushveld gerbil (Tatera spp.) 4

Pouched mouse (Saccostomus spp.) 14

Large-eared mouse (Malacothrix spp.) 1

Dormouse (Graphiurus spp.) 5

Mole-rat (Cryptomys spp.) 4

Crested porcupine (Hystrix cristata) 3

Hyracoidea

Rock hyrax (Procavia capensis) 172

Eulipotyphla

Southern African hedgehog (Atelerix frontalis) 3

Lesser red musk shrew (Crocidura hirta) 3

Macroscelidea

Round-eared sengi (Macroscelides proboscideus) 3

Bushveld elephant shrew (Elephantulus intufi) 6

Chiroptera

Slit-faced Bat (Nycteris spp.) 1

Horseshoe bat (Rhinolophus spp.) 1

Egyptian free-tailed bat (Tadarida aegyptiaca) 2

Leaf-nosed bat (Hipposideros spp.) 2

Primates

Chacma baboon (Papio ursinus) 9

Tubulidentata

Aardvark (Orycteropus afer) 2

Aves

Galliformes 74

Helmeted guineafowl (Numida meleagris) 35

Francolin (Pternistes spp.) 13

Pteroclidiformes

Sandgrouse (Pterocles spp.) 5

Charadriiformes

Kurrichane buttonquail (Turnix sylvaticus) 3

Crowned lapwing (Vanellus coronatus) 1

Spotted thick-knee (Burhinus capensis) 10

Anseriformes

Egyptian goose (Alopochen aegyptiaca) 1

Duck (Anas spp.) 1

Gruiformes

Red-knobbed coot (Fulica cristata) 1

Otidiformes

Kori bustard (Ardeotis kori) 6

Ludwig's bustard (Neotis ludwigii) 6

Karoo korhaan (Eupodotis vigorsii) 8

Red-crested korhaan (Lophotis ruficrista) 7

Northern black korhaan (Afrotis afraoides) 23

Struthioniformes

Ostrich (Struthio camelus) 1

Strigiformes

Barn owl (Tyto alba) 8

Spotted eagle-owl (Bubo africanus) 2

Verreaux's eagle-owl (Bubo lacteus) 1

Accipitriformes

Pale chanting goshawk (Melierax canorus) 1

Brown snake eagle (Circaetus cinereus) 1

Columbiformes

Cape turtle-dove (Streptopelia capicola) 5

Laughing dove (Spilopelia senegalensis) 4

Namaqua dove (Oena capensis) 2

African green pigeon (Treron calvus) 1

Bucerotiformes Southern yellow-billed hornbill (Tockus leucomelas) 1

Coraciiformes

Purple roller (Coracias naevius) 2 75

Caprimulgiformes

Nightjar (Caprimulgus spp.) 9

Passeriformes

Fork-tailed drongo (Dicrurus adsimilis) 1

Lark (Mirafra spp.) 2

Groundscraper thrush (Psophocichla litsitsirupa) 2

Ant-eating chat (Myrmecocichla formicivora) 1

Pipit (Anthus spp.) 3

Starling (Lamprotornis spp.) 1

Long-tailed paradise whydah (Vidua paradisaea) 6

Shaft-tailed whydah (Vidua regia) 1

Sociable weaver (Philetairus socius) 3

Bunting (Emberiza spp.) 2

Amphibia

Anura

African bullfrog (Pyxicephalus adspersus) 5

Reptilia

Squamata

Puff adder (Bitis arietans) 5

Cape cobra (Naja nivea) 2

Brown house snake (Boaedon spp.) 1

Egg-eating snake (Dasypeltis spp.) 1

Mole snake (Pseudaspis cana) 2

African rock python (Python sebae) 1

Rock monitor (Varanus albigularis) 9

Agama lizard (Agama spp.) 5

Flap-necked chameleon (Chamaeleo dilepis) 2

Namaqua chameleon (Chamaeleo namquensis) 1

Gecko spp. 5

Giant Ground Gecko (Chondrodactylus angulifer) 5

Skink (Trachylepis spp.) 3

Testudines

African side-necked turtle (Pelomedusa subrufa) 2

FIGURE LEGENDS

Figure 1.-- Study area for southern Namibian caracal (Caracal caracal) dietary study with habitat delineations for Karoo/Karasberg, Kalahari grassland, sparse shrubland and Acacia thornbrush habitats.

TABLES

Table 1.-- Total prey items in the diets of southern Namibian caracals (Caracal caracal) as determined by kill site, stomach content and scat analyses from 2010-2016.

Contribution of All Prey Total Prey Items Items n = 1381 (relative frequency) Mammalia 1149 83.2 Artiodactyla 544 39.4 Carnivora 81 5.9 Chiroptera 6 0.4 Eulipotyphla 6 0.4 Hyracoidea 172 12.5 Lagomorpha 123 8.9 Macroscelidea 9 0.7 Primates 9 0.7 Rodentia 168 12.2 Tubulidentata 2 0.1

Domestic stock 29 2.1

Aves 183 13.3 Accipitriformes 2 0.1 Anseriformes 3 0.2 Bucerotiformes 1 0.1 Caprimulgiformes 9 0.7 Charadriiformes 14 1.0 Columbiformes 13 0.9 Coraciiformes 2 0.1 Galliformes 48 3.5 Gruiformes 1 0.1 Otidiformes 50 3.6 Passeriformes 22 1.6 Pteroclidiformes 5 0.4 Strigiformes 12 0.9 Struthioniformes 1 0.1

Amphibia 5 0.4 Anura 5 0.4

Reptilia 44 3.2 Squamata 42 3.0 Testudines 2 0.1

Table 2.-- Relative frequencies of prey items in the diets of southern Namibian caracals (Caracal caracal) from Karoo/Karasberg (KK), Kalahari grassland (KG), sparse shrubland (SS) and Acacia thornbrush (AT) habitats from 2010-2016.

Prey Type All (N=1140) KK (n=324) KG (n=473) SS (n=203) AT (n=140) Mammalia 83.2 89.7 89.2 80.2 62.1 Artiodactyla 39.4 23.7 64.3 38.8 9.4 Carnivora 5.9 4.2 8.5 7.2 1.3 Chiroptera 0.4 0.8 0.2 0.0 0.9 Eulipotyphla 0.4 0.0 0.8 0.4 0.4 Hyracoidea 12.5 43.4 0.0 0.0 3.0 Lagomorpha 8.9 6.3 5.3 16.9 13.2 Macroscelidea 0.7 0.0 0.4 1.3 1.7 Primates 0.7 2.4 0.0 0.0 0.0 Rodentia 12.2 7.6 7.6 10.5 31.5 Tubulidentata 0.1 0.0 0.4 0.0 0.0 Domestic stock 2.1 1.3 1.9 5.1 0.9 Aves 13.3 7.9 9.3 13.1 31.1 Accipitriformes 0.1 0.0 0.0 0.4 0.4 Anseriformes 0.2 0.0 0.0 0.0 1.3 Bucerotiformes 0.1 0.0 0.0 0.0 0.4 Caprimulgiformes 0.7 0.5 0.2 1.3 1.3 Charadriiformes 1.0 0.0 1.5 0.4 2.1 Columbiformes 0.9 0.5 0.8 0.8 2.1 Coraciiformes 0.1 0.0 0.0 0.0 0.9 Galliformes 3.5 0.5 3.6 0.8 10.6 Gruiformes 0.1 0.0 0.0 0.0 0.4 Otidiformes 3.6 3.7 1.7 6.8 4.7 Passeriformes 1.6 1.1 0.6 1.3 5.1 Pteroclidiformes 0.4 0.0 0.8 0.4 0.0 Strigiformes 0.9 1.3 0.2 0.8 1.7 Struthioniformes 0.1 0.3 0.0 0.0 0.0 Amphibia 0.4 0.0 0.0 0.0 2.1 Anura 0.4 0.0 0.0 0.0 2.1 Reptilia 3.2 2.4 1.5 6.8 4.7 Squamata 3.0 2.4 1.5 6.8 3.8 Testudines 0.1 0.0 0.0 0.0 0.9

Table 3.-- Relative frequencies of prey items in the diets of southern Namibian caracals (Caracal caracal) represented by three different methods including stomach contents, kill sites and scat samples from 2010-2016.

Stomachs Kill sites Scats n = 202 688 250 Total recorded prey: 328 688 365 Mammalia 74.1 90.6 77.5 Artiodactyla 26.2 53.6 24.4 Carnivora 4.0 7.8 3.8 Chiroptera 0.9 0.0 0.8 Eulipotyphla 0.6 0.3 0.5 Hyracoidea 9.5 15.8 8.8 Lagomorpha 8.8 7.1 12.3 Macroscelidea 0.9 0.0 1.6 Primates 0.6 0.6 0.8 Rodentia 18.0 3.8 22.7 Tubulidentata 0.6 0.0 0.0

Domestic stock 4.0 1.5 1.6

Aves 20.1 7.6 17.8 Accipitriformes 0.0 0.3 0.0 Anseriformes 0.6 0.0 0.3 Bucerotiformes 0.0 0.1 0.0 Caprimulgiformes 0.9 0.1 1.4 Charadriiformes 1.8 0.6 1.1 Columbiformes 2.4 0.1 1.1 Coraciiformes 0.3 0.0 0.3 Galliformes 3.7 2.5 5.2 Gruiformes 0.3 0.0 0.0 Otidiformes 4.6 2.5 4.9 Passeriformes 4.0 0.3 1.9 Pteroclidiformes 0.6 0.1 0.5 Strigiformes 0.6 0.9 1.1 Struthioniformes 0.3 0.0 0.0

Amphibia 0.9 0.0 0.5 Anura 0.9 0.0 0.5

Reptilia 4.9 1.9 4.1 Squamata 4.6 1.9 3.8 Testudines 0.3 0.0 0.3

Table 4.-- Relative frequencies of prey items in the diets of southern Namibian caracals (Caracal caracal) in Karoo/ Karasberg habitat as represented by three different methods including stomach contents, kill sites and scat samples from 2010-2016.

Prey Type Total Stomachs Kill sites Scats Mammalia 89.7 84.3 94.5 82.3 Artiodactyla 23.7 18.1 28.0 17.7 Carnivora 4.2 1.2 6.4 1.3 Chiroptera 0.8 1.2 0.0 2.5 Eulipotyphla 0.0 0.0 0.0 0.0 Hyracoidea 43.4 36.1 50.0 32.9 Lagomorpha 6.3 6.0 5.0 10.1 Macroscelidea 0.0 0.0 0.0 0.0 Primates 2.4 2.4 1.8 3.8 Rodentia 7.6 15.7 2.8 12.7 Tubulidentata 0.0 0.0 0.0 0.0 Domestic stock 1.3 3.6 0.5 1.3 Aves 7.9 10.8 5.0 12.7 Accipitriformes 0.0 0.0 0.0 0.0 Anseriformes 0.0 0.0 0.0 0.0 Bucerotiformes 0.0 0.0 0.0 0.0 Caprimulgiformes 0.5 0.0 0.5 1.3 Charadriiformes 0.0 0.0 0.0 0.0 Columbiformes 0.5 1.2 0.0 1.3 Coraciiformes 0.0 0.0 0.0 0.0 Galliformes 0.5 0.0 0.5 1.3 Gruiformes 0.0 0.0 0.0 0.0 Otidiformes 3.7 6.0 2.3 5.1 Passeriformes 1.1 1.2 0.5 2.5 Pteroclidiformes 0.0 0.0 0.0 0.0 Strigiformes 1.3 1.2 1.4 1.3 Struthioniformes 0.3 1.2 0.0 0.0 Amphibia 0.0 0.0 0.0 0.0 Anura 0.0 0.0 0.0 0.0 Reptilia 2.4 4.8 0.5 5.1 Squamata 2.4 4.8 0.5 5.1 Testudines 0.0 0.0 0.0 0.0

Table 5.-- Relative frequencies of prey items in the diets of southern Namibian caracals (Caracal caracal) in Kalahari grassland habitat as represented by three different methods including stomach contents, kill sites and scat samples from 2010-2016.

Prey Type Total Stomachs Kill sites Scats Mammalia 89.2 81.4 93.2 83.3 Artiodactyla 64.3 45.1 74.8 46.7 Carnivora 8.5 6.9 8.9 8.9 Chiroptera 0.2 1.0 0.0 0.0 Eulipotyphla 0.8 2.0 0.3 1.1 Hyracoidea 0.0 0.0 0.0 0.0 Lagomorpha 5.3 5.9 4.5 7.8 Macroscelidea 0.4 1.0 0.0 1.1 Primates 0.0 0.0 0.0 0.0 Rodentia 7.6 13.7 3.6 15.6 Tubulidentata 0.4 2.0 0.0 0.0 Domestic stock 1.9 3.9 1.2 2.2 Aves 9.3 14.7 6.5 13.3 Accipitriformes 0.0 0.0 0.0 0.0 Anseriformes 0.0 0.0 0.0 0.0 Bucerotiformes 0.0 0.0 0.0 0.0 Caprimulgiformes 0.2 0.0 0.0 1.1 Charadriiformes 1.5 3.9 0.6 2.2 Columbiformes 0.8 2.0 0.3 1.1 Coraciiformes 0.0 0.0 0.0 0.0 Galliformes 3.6 2.9 3.3 5.6 Gruiformes 0.0 0.0 0.0 0.0 Otidiformes 1.7 1.0 1.8 2.2 Passeriformes 0.6 2.9 0.0 0.0 Pteroclidiformes 0.8 2.0 0.3 1.1 Strigiformes 0.2 0.0 0.3 0.0 Struthioniformes 0.0 0.0 0.0 0.0 Amphibia 0.0 0.0 0.0 0.0 Anura 0.0 0.0 0.0 0.0 Reptilia 1.5 3.9 0.3 3.3 Squamata 1.5 3.9 0.3 3.3 Testudines 0.0 0.0 0.0 0.0

Table 6.-- Relative frequencies of prey items in the diets of southern Namibian caracals (Caracal caracal) in sparse shrubland habitat as represented by three different methods including stomach contents, kill sites and scat samples from 2010-2016.

Prey Type Total Stomachs Kill sites Scats Mammalia 80.2 75.8 83.3 78.7 Artiodactyla 38.8 29.0 46.5 34.4 Carnivora 7.2 6.5 8.8 4.9 Chiroptera 0.0 0.0 0.0 0.0 Eulipotyphla 0.4 0.0 0.9 0.0 Hyracoidea 0.0 0.0 0.0 0.0 Lagomorpha 16.9 14.5 17.5 18.0 Macroscelidea 1.3 1.6 0.0 3.3 Primates 0.0 0.0 0.0 0.0 Rodentia 10.5 16.1 5.3 14.8 Tubulidentata 0.0 0.0 0.0 0.0 Domestic stock 5.1 8.1 4.4 3.3 Aves 13.1 17.7 7.9 18.0 Accipitriformes 0.4 0.0 0.9 0.0 Anseriformes 0.0 0.0 0.0 0.0 Bucerotiformes 0.0 0.0 0.0 0.0 Caprimulgiformes 1.3 3.2 0.0 1.6 Charadriiformes 0.4 0.0 0.9 0.0 Columbiformes 0.8 3.2 0.0 0.0 Coraciiformes 0.0 0.0 0.0 0.0 Galliformes 0.8 0.0 0.0 3.3 Gruiformes 0.0 0.0 0.0 0.0 Otidiformes 6.8 9.7 4.4 8.2 Passeriformes 1.3 1.6 0.9 1.6 Pteroclidiformes 0.4 0.0 0.0 1.6 Strigiformes 0.8 0.0 0.9 1.6 Struthioniformes 0.0 0.0 0.0 0.0 Amphibia 0.0 0.0 0.0 0.0 Anura 0.0 0.0 0.0 0.0 Reptilia 6.8 6.5 8.8 3.3 Squamata 6.8 6.5 8.8 3.3 Testudines 0.0 0.0 0.0 0.0

Table 7.-- Relative frequencies of prey items in the diets of southern Namibian caracals (Caracal caracal) in Acacia thornbrush habitat as represented by three different methods including stomach contents, kill sites and scat samples from 2010-2016.

Prey Type Total Stomachs Kill sites Scats Mammalia 62.1 53.1 42.1 70.4 Artiodactyla 9.4 8.6 15.8 8.9 Carnivora 1.3 1.2 0.0 1.5 Chiroptera 0.9 1.2 0.0 0.7 Eulipotyphla 0.4 0.0 0.0 0.7 Hyracoidea 3.0 1.2 0.0 4.4 Lagomorpha 13.2 11.1 15.8 14.1 Macroscelidea 1.7 1.2 0.0 2.2 Primates 0.0 0.0 0.0 0.0 Rodentia 31.5 27.2 10.5 37.0 Tubulidentata 0.0 0.0 0.0 0.0 Domestic stock 0.9 1.2 0.0 0.7 Aves 31.1 38.3 52.6 23.7 Accipitriformes 0.4 0.0 5.3 0.0 Anseriformes 1.3 2.5 0.0 0.7 Bucerotiformes 0.4 0.0 5.3 0.0 Caprimulgiformes 1.3 1.2 0.0 1.5 Charadriiformes 2.1 2.5 5.3 1.5 Columbiformes 2.1 3.7 0.0 1.5 Coraciiformes 0.9 1.2 0.0 0.7 Galliformes 10.6 11.1 26.3 8.1 Gruiformes 0.4 1.2 0.0 0.0 Otidiformes 4.7 3.7 5.3 5.2 Passeriformes 5.1 9.9 0.0 3.0 Pteroclidiformes 0.0 0.0 0.0 0.0 Strigiformes 1.7 1.2 5.3 1.5 Struthioniformes 0.0 0.0 0.0 0.0 Amphibia 2.1 3.7 0.0 1.5 Anura 2.1 3.7 0.0 1.5 Reptilia 4.7 4.9 5.3 4.4 Squamata 3.8 3.7 5.3 3.7 Testudines 0.9 1.2 0.0 0.7

FIGURES

APPENDIX B: SURVIVAL, MORTALITY & REPRODUCTION OF CARACALS (CARACAL CARACAL) ON NAMIBIAN FARMLANDS

Aletris M. Neils

(In the format of Biological Conservation)

Survival, mortality & reproduction of caracals (Caracal caracal) on Namibian farmlands Running head: Caracal survival on Namibian farmlands

Article impact statement: Life history schedules and survival analysis for Namibian caracals reveal stock farms are a population sink due to anthropogenic mortality.

Keywords: Life table, survival curve, Caracal caracal, human-wildlife conflict

Word count: 4323

Author: Aletris M. Neils, [email protected]; Conservation CATalyst, 3755 West Driscol Lane, Tucson, Arizona 85745

Acknowledgements

First and foremost, the author would like to express gratitude to local farming hosts,

Kiripotib Guest Farm (Hans & Claudia von Hase), Timm & Inez Miller, and Tyse & Annetta

Swart. I also acknowledge the many livestock farmers that contributed specimens and information; I could not have done it without your cooperation and trust. Field research was conducted with meticulous detail in part by the exceptional Elin Gromberg and Magnus Nystrand.

I am very grateful for field assistance from several passionate interns especially Elan Fox, Liliy

Harrison, and many others who contributed to the success of the project through the years.

Necropsy protocols were developed with direction and guidance from Dr. Axel Hartmann. I thank

John Koprowski for his guidance and multiple reviewers for constructive comments. This research was conducted with support from the Namibian Ministry of Environment and Tourism (permit number 1539). Funding was provided by Fulbright, Cheetah Kids, Crosby Family, B. Kay, M.

Sloan, and contributions from many other donors.

SURVIVAL, MORTALITY & REPRODUCTION OF CARACALS (CARACAL CARACAL) ON

NAMIBIAN FARMLANDS

ABSTRACT

Caracals (Caracal caracal) are a highly persecuted species in southern Namibia due to perceived threats to small stock. Caracal persecution has occurred on a large scale for decades, but the species appears to persist while many larger carnivores have been effectively extirpated from the region. Future population collapse can be predicted by examining population demographics and age distributions. Data were collected from radio-collared and necropsied caracals from 2010-

2016 to estimate key demographic parameters including survival and reproduction values. These data were used to generate life history schedules and Kaplan-Meier survival curves to determine whether caracal populations on southern Namibian farmlands can withstand such high anthropogenic mortality. Life table data revealed a net reproductive rate >1.00 for the population but only when an even sex ratio of kittens was assumed. Southern Namibia stock farms probably represent a population sink for caracals, whereas the local population relies on immigration from adjacent areas in order to persist. Although immigration in heavily hunted felid populations compensates for high losses, it also leads to a demographic shift towards younger animals as observed in this study resulting in population instability under the influence of high male turnover.

Caracal populations are projected to decline if persecution pressure continues.

INTRODUCTION

Human–wildlife conﬂict (HWC) arises when a species poses a direct and recurring threat to community livelihood. Rangelands have always been particularly prone to HWC; this is often where carnivores, prey, people and livestock occur together in the same space and are likely to encounter one another (Zimmerman 2010). HWC such as livestock depredation and subsequent predator control responses by farmers are a growing concern worldwide (Loveridge 2010).

Livestock depredation often results in persecution efforts of predators, defined as the persistent killing, chasing or other harassment of a species with the goal of eradication, and includes both retaliatory and preventative killings (Zimmerman et al. 2010). Predator numbers are in decline globally in part due to these conflicts (Loveridge et al. 2010).

Carnivore persecution in southern Africa has occurred on a large scale for decades (Stuart

& Hickman 1991; Stuart & Stuart 2013). In southern Namibia, large carnivores such as lion

(Panthera leo), African wild dog (Lycaon pictus), and spotted hyena (Crocuta crocuta) have been systematically extirpated due to persecution from farmers following European settlement. Only fragmented remnant populations of leopard (Panthera pardus) and cheetah (Acinonyx jubatus) occur. Mesocarnivores such as caracals (Caracal caracal) and black-backed jackals (Canis mesomelas) still remain but are now considered the greatest predatory threat to small stock such as sheep and goats by local farmers (Stuart & Hickman 1991; Mukherjee et al. 2004; Stein 2006).

Caracals are a poorly understood wild felid that have had little research attention. Little has been published of their basic ecology, behavior, and life history traits and sample sizes have been small (Marker & Dickman 2005; Sunquist & Sunquist 2002). No science-based population estimates exist for caracals even though caracals are considered threatened, endangered or on the 89

verge of extinction throughout a majority of their range (Sunquist & Sunquist 2002). In sub-

Saharan Africa, caracals are protected in about half of the countries in their range (Nowell &

Jackson 1996), but caracal population size and densities in southern Africa are rough estimates based on very little scientific information. Caracals receive no legal protection or conservation actions in Namibia, one of the last remaining geographic strongholds for the species (Sunquist &

Sunquist 2002).

Caracals have long been considered the greatest predatory threat to small stock such as sheep and goats by farmers in Namibia (Stuart & Hickman 1991; Mukherjee et al. 2004).

Namibian farmers consider caracals one of the top threats to small stock (Stein 2006) and may legally remove caracals without restriction. Prior to the 1990s, most research programs on caracals consisted of finding effective means of exterminating the species. From 1931-1952, Stuart (1984) asserted that over 2,200 caracals per year were killed in control operations in the Karoo, South

Africa. Similarly, farmers in Namibia reported killing up to 2,800 caracals in 1981 alone and much killing likely goes unreported (Nowell & Jackson 1996). Records on numbers of caracals killed each year are probably gross underestimations since caracal pelts are of little commercial value

(Sunquist & Sunquist 2002). Persecution has lasted for decades on a regional scale and continues to this day (Stuart & Stuart 2013).

Caracals have appeared to so far withstand rigorous eradication campaigns in South Africa and Namibia. Caracal populations have been claimed to be unaffected by persecution and eradication campaigns and were thought to have the capacity to withstand indefinite persecution pressure (Stuart 1981; Stuart 1986). However, eventual long-range declines were also predicted

(Visser 1976). At present, caracal numbers may be dwindling and a current population assessment 90

is needed to determine if declines have occurred. Effects of caracal persecution are unknown, but in other carnivore populations these include altered population dynamics (Sacks 2005; Cooley et al. 2009), increased infanticide (Stuart 1981; Swenson et al. 1997; Cooley et al. 2009), and decreased genetic diversity due to a reduced effective population size (Creel 1998). Since carnivores often serve as top-down ecosystem regulators, potential ecosystem effects of carnivore removal include overpopulation of prey species and increased disease among prey populations

(Woodroffe & Ginsberg 2005).

Conservation and management of species experiencing heavy anthropogenic impacts can greatly benefit from detailed demographic information; future population collapse can be predicted by examining population demographics and age distributions (Caughley 1977; White 2000). By examining vital statistics of caracals on Namibian farmlands the status and population trends of caracals can be assessed. Such data have never been collected for caracals.

METHODS

2.1. Study area

Southern Namibia is one of the last strongholds of caracals in the world (Sunquist &

Sunquist 2002). This research was conducted on privately owned livestock farmlands in southern

Namibia, from the village of Dordabis (70km southwest of Windhoek) south to the Groot Karas

Mountains (130km southwest of Keetmanshoop). Research was conducted in three study sites representing distinct habitat types in southern Namibia (Karoo/ Karasberg, Kalahari grasslands, and sparse shrublands). The region is extremely arid, with highly fluctuating and unpredictable

rainfall (Strohbach 2011). Large tracts of grazing habitat are relatively scarce and insufficient to support large scale cattle farming; therefore, commercial farmers rely on small stock, primarily sheep and goats (Quan et al. 2004). To a lesser extent, wild game such as kudu (Tragelaphus strepsiceros), oryx (Oryx gazelle) and springbok (Antidorcas marsupialis) are used by landowners for subsistence and commercial hunting. Large carnivores such as lion, leopard, cheetah and spotted hyena have already been extirpated due to persecution from ranchers.

2.2. Animal trapping, handling, and immobilization

Study animals used for this research included live-trapped and killed caracals captured and donated by farmers. Killed caracals included fresh, frozen, and desiccated specimens. An accurately represented cross-section of the population was assumed for this study due to the variety of methods used by southern Namibian farmers to kill caracals. These include shooting at night by spotlight (with and without predator- calling), steel gin traps/ leg-hold traps, cage traps, neck or body snares, jackal canons (a wire-triggered gun typically rigged to fences and burrows), poisons, hunting dogs, and deliberate vehicular collision. Handling of animals followed the

American Society of Mammalogist’s Animal Care and Use Committee protocol

(http://www.mammalsociety.org/committees/animal-care-and-use). The project was conducted under University of Arizona’s IACUC certification number 6809 and under research permit 1539 granted by the Namibian Ministry of the Environment.

remote injection gun was used to immobilize cats opportunistically caught in chicken coops, livestock pens, snares or leghold traps. During immobilization, cats were placed on their left side and monitored continuously.

Sedated caracals were examined for general health and breeding condition, morphological data (lengths and body mass) were collected and animals were aged by tooth coloration and wear

(Stuart & Stuart 1985; Stander 1997). Based on examinations of teeth, caracals were categorized into age cohorts for analysis (Stuart & Stuart 1985; Stander 1997). VHF and GPS radio collars

(Followit, Tellus GPS; Lindesburg, Sweden) were fitted around the neck of tagged cats. Collars weighed between 285-370 g (did not exceed 3% of the animal’s weight) and were equipped with mortality sensors to allow a timely response to find deceased study animals. Cats were released within 500 m of capture location.

2.3. Monitoring

Following release, radio-collared cats were monitored every three hours via GPS collars

At least once per week, cats were triangulated and approached on foot to within 1km, where store- on-board GPS points were downloaded using UHF. Mortality signals were investigated immediately. All GPS collars were labeled with contact information caracal deaths could be reported by farmers. Necropsies were conducted within 24 hours of receipt of carcasses.

2.4. Survival and reproduction

Data from radio collared and necropsied caracals were used to determine mortality and reproductive rates for the population. In addition to examining anesthetized caracals prior to radio collaring, necropsies were performed on all available deceased caracals to determine age and cause of death. All available carcasses of female caracals were examined for signs of proestrus and pregnancy (lactation, corpora lutea and/or embryos). Placental scars from necropsied caracals were used to determine past reproductive history based on methodology and criteria described by

Mowat et al. (1996). By monitoring collared reproductive females, dens were located and camera- traps were used to document litter sizes and determine age-specific fecundity.

Ages were determined with precision for study animals under one year of age and for animals in their second year of age based on morphological characteristics (Stuart & Stuart 1985).

Based primarily on examinations of teeth, as well as reproductive history and other physical features, adult caracals were categorized into annual age cohorts >2 years of age for analysis

(Stuart & Stuart 1985; Stander 1997). Disappearance or removal from the population was considered to be equivalent to mortality, as some injured caracals were placed in captivity rather than being immediately euthanized by the farmer.

2.5. Life tables

Mortality and fecundity schedules are powerful tools to analyze demographic trends and vital statistics of wild populations. Time-specific life tables represent an assemblage of a population’s life history data organized by age cohorts and life table statistics are used to estimate

rates of population change (Caughley 1966). Using all caracals (radio-collared and necropsied), time-specific cohort life tables with staggered entry were constructed for the caracal population across all study areas. For all caracals (pooled male and female) and for male caracals alone, number alive (nx) was calculated for each age class, as was survivorship (lx; nx / n0), number dying

(dx; nx - nx+1), death rate (qx; dx / nx), survival rate (1 – qx), and estimated number of age classes

(years) left to be lived (Tx) (Caughley 1966). For female-only life table schedules, birth rate

(average number of female young born to breeding females, bx) was also calculated and used to determine net reproductive rate (r0), or the average number of daughters born to a female if she passed through her lifetime conforming to the calculated mortality rates calculated in the life table schedule (Caughley 1966). Fates (mortality) were known definitively for each study animal.

Composite cohort life tables were constructed for males and females, first captured as adults or post-weaned sub-adults, which were assigned to different age cohorts based on age estimations

(described in Section 2.6). All caracals aged and included in the life table schedules were of weaned age, although some were still travelling with their mother. By including medium-brown placental scars, reproductive estimates were conservative, as light-to-medium scars may also result from re-absorption of embryos, i.e. embryos that were never born (Mowat et al. 1996). Counting placental scars to define reproduction assumed embryos survived to birth (Mowat et al. 1996).

This study assumed an even sex ratio of male: female at birth. A stable age distribution was not assumed for this analysis.

2.6. Incorporating estimated age and reproduction into life tables

All caracals first obtained as adults (both captured and necropsied) and were aged to closest estimated year according to criteria from Stuart & Stuart (1985) and Stander (1997). Age estimates primarily included examination of dentition and enamel wear. Cementum annuli cross-sections and examination of premolar wear (Trainer & Matson 1988) were conducted for a random subset of caracals to refine and confirm aging accuracy. Although accuracy of aging caracals by visual examination is increasing difficult after 3-4 years (Stander 1997), most study animals (93%) did not exceed this age class due to human persecution. Thus, increasingly inaccurate age estimations based on old age were not a significant factor in this study. Caracals >5 years of age were counted as a single cohort, but this cohort was only reached by 4 individuals (2%). Only a single caracal examined might have approached the reported life expectancy of the species (determined by extreme tooth wear).

Placental scars, presence of embryos, and signs of lactation were used to determine caracal reproduction for life tables using methods described by Mowat et al. (1996). Females observed to be either lactating or post-lactation and/or exhibited medium-to-dark placental scars were considered reproductive in the current age class. Non-lactating females with light brown placental scars were considered to be reproductive the previous year. The presence of embryos determined female reproductive status based on the estimated date of birth.

2.7. Survival curves

Staggered entry Kaplan-Meier survival curves were generated using MedCalc

(www.medcalc.org) for radio-collared caracals to compare survival rates of different caracal age 96

classed. Associated p-values were used to compare differences between habitat types, sex, and age. Curves were generated for all male and female caracals (pooled habitats), all adults and yearling/ sub-adults (pooled habitats), and all caracals by habitat type (pooled age and sex). Log- rank tests (chi- squared) were used to compare survival distributions of different groups (Peto &

Peto 1972).

RESULTS

3.1. Captures and necropsies

Data from 208 caracals were collected throughout the course of this project, 34 radio- collared caracals (41% female, 59% male) and 174 necropsied caracals (45% female, 55% male)

(Table 1). Collared caracals showed a different age distribution according to sex (36% yearling or sub-adult females and 64% adult females vs. 75% yearling or sub-adult males and 25% adult males) (Table 1). The sex of one caracal could not be determined due to extreme trauma and this animal was only included in age analyses. Necropsied caracals showed a more similar age distribution according to sex (58% yearling or sub-adult females; 42% adult females and 69% yearling or sub-adult males and 31% adult males) (Table 1). Although most of the study animals were acquired only after having been killed by humans, all radio- collared caracals in this study met the same fate.

3.2. Life tables

The majority of post-weaning caracals (84%) were projected to survive to the end of their first year (Table 2). Only 36% were projected to survive to the end of their second year, 16% to the end of their third year, and <7% were expected to survive to the end of their fourth year (Table

2). Caracals only had a ~2% chance of surviving past their 5th birthday (Table 2). Only one individual male could have significantly exceeded this age based on extreme dental wear. Female caracals experienced higher survivorship between years 2-3 compared to males (~45% vs ~30%)

(Table 3). This may be a product of reproductive behavior where females of this age are denning rather than travelling vast distances like males, and are less exposed to anthropogenic hazards. All cats demonstrated a projected 40-46% chance of survival each year for years 2-4, after which the chances dropped precipitously (Table 2). Life expectancy for all caracals was by far the highest in the first year (Tables 2-5).

3.4. Survival curves

Kaplan-Meier curves for radio-collared caracal factored by sex (M-F) were different; male caracals experienced lower survival probabilities than females (chi-squared= 3.6152; P = 0.057;

Fig. 1). Survivorship curves factored by age (yearling/ subadult vs. adults) were different with younger animals experiencing lower survival (chi-squared= 13.4228; P = 0.0002; Fig. 2).

Survivorship curves did not differ by habitat type (chi-squared= 0.07851; P = 0.96; Fig. 3).

Summary statistics for sex, age, and habitat type are provided in Table 6.

3.5. Reproduction

Reproductive rates were highest for 4-5 year old caracals (1.58; Table 4) followed by 3-4 year old caracals (1.34; Table 4). Caracals 2-3 years old averaged just over one female offspring

(Table 4). Reproductive rates declined after 5 years (1.25; Table 4). In addition to life table statistics calculated for female caracals assuming an even sex ratio of kittens, a second life table was constructed for female caracals assuming a female: male ratio of 45: 55 for kittens; the observed sex ratio for all deceased study animals. An accurately represented cross-section of the population was assumed for deceased study animals due to the variety of methods used by southern

Namibian farmers to kill caracals (see Methods 2.2). A net reproductive rate (r0) of 1.06 was determined assuming an even sex ratio of kittens (Table 4), but fell to 0.891 assuming a male- skewed ratio as was observed in the field for adults and subadult caracals (Table 5).

Low reproductive output of second year individuals may be due to inherently lower reproductive success of young mothers (Smith 2009). Litter sizes averaged 2.2 (+1.09) from the second year onward. On only two occasions were females documented with four kittens. Both of these females had survived to higher age classes and were both exceptionally large; one of these females was over 12 kg. Camera traps placed at three different den sites recorded single kittens twice, and twins once. Both single kittens belonged to 3-4 year old females, below the average estimated number of young for that cohort.

DISCUSSION

Caracals are persecuted primarily due to perceived threats to livestock. Caracal mortality stems principally from anthropogenic forces, as eradication of caracals is fully legal in Namibia 99

based on their current classification as ‘vermin’ (Stuart and Hickman 1991; Mukherjee et al. 2004).

All caracals in this study were killed by humans. Southern Namibian caracals experience low adult survival and display non-typical life expectancies indicating a population under intense hunting pressure (Sibly et al. 1997).

4.1. Felid response to high mortality

Depending on assumed caracal sex ratio at birth, net reproductive rates from life table schedules ranged from 0.891- 1.06, suggesting that caracals may not be able to withstand current rates of human persecution. Based on these findings, a population decline would be expected in the absence of immigration from source populations. Resilience of caracal populations likely depends on high levels of sub-adult immigration from neighboring areas and philopatric recruitment of female offspring, as is observed in heavily hunted puma populations in the United

States (Lindzey et al. 1992, Sweanor et al. 2000). Increased immigration of young males following resident removal has been observed for bobcats (Lynx rufus), pumas (Puma concolor) and African lions (Rolley 1985; Ross & Jalkotsky 1992; Slough & Mowat 1996; Sweanor et al. 2000; Kamler et al. 2000; Lambert et al 2006; Cooley et al. 2009; Barthold et al. 2016). Frequent deaths of adult male pumas and African lions are known to disrupt social structure of the population (Sweanor et al. 2000; Barthold et al. 2016). Increased immigration and recruitment of young animals from adjacent areas results in little or no change in densities, but shifts population structure towards young animals (Robinson et al. 2008). The southern Namibian caracal population may be experiencing constant influx and rapid turnover of young males.

100

In heavily hunted populations with low adult survival, life expectancy is highest in year 0-

1 (Mosby 1969; Sibly et al. 1997). While long distance dispersal (especially of males) compensates for high losses within a population, ecological consequences exist. High male turnover leads to instability of the population, decreased resilience, declining female: male ratios, and increased infanticide (Sweanor et al 2000; Cooley et al. 2009; Wielgus et al 2013; Barthold et al 2016). Populations composed of young and inexperienced individuals are more susceptible to variable habitat and environmental conditions and stochastic events such as drought (Gordon et al.

2004). To compound this issue, farmer persecution efforts typically increase during droughts

(unpulished data). Ironically, destroying caracals may increase conflicts by creating a vacuum constantly filled by young immigrants. As the population structure shifts toward young animals, the management goal of reduced conﬂict may be negated as young felids are most often involved in conﬂicts with humans (Beier 1991).

4.2. Population sinks

Many carnivores display density-independent long dispersal by male individuals and such immigration strategies can increase local population densities regardless of birth and death rates

(Franke and Woodroffe 2001). Caracal removal in southern Namibian may result in little or no reduction in local densities but can shift population structure towards young animals. Hunting pressure that is uneven across a broad landscape is known to induce source-sink dynamics in carnivore populations (Mowat 1996). Attractive habitat patches that provide quality resources may also exhibit disparate rates of anthropogenic mortality that cannot be immediately perceived by immigrants (Delibes et al. 2001). Habitat that fits these criteria can become a population sink, 101

where a continual flow of young animals arrive into quality habitat yet experience high mortality

(Pulliam 1988, Thomas and Kunin 1999). Namibian farms appear to represent a mosaic of population sinks for caracals and the population likely depends on immigration from adjacent areas to persist.

4.3. Caracal kitten sex ratios and survival

Net reproductive rates for this study were conservative as kitten survival to birth was assumed at 100%; it is possible that not all kittens conceived survived gestation. Slight shifts in female: male sex ratio of kittens determined whether estimated net reproductive rates fell above or below 1.00, indicating a growing or shrinking population respectively. Documenting sex ratios of caracal kittens at birth is a priority, as some mammals appear to have an innate ability to alter sex- ratios of young produced under certain environmental conditions (Clutton-Brock & Iason 1986).

Mammals that experience high male mortality may skew sex ratios at birth towards males

(Cockburn et al. 2002), and in some carnivores younger mothers will overproduce male offspring

(Cockburn et al. 2002). Sex ratio of yearling caracals in this study was strongly skewed towards males. If young caracal mothers indeed overproduce male offspring, estimated net reproductive rate for this population would decline.

Caracal kitten survival to weaning was unknown but is likely much lower than 100% in the study area. Besides male conspecifics, caracal kittens have been documented being killed by a variety of predators including leopards, cheetahs, hyenas, black-backed jackals (Canis mesomelas), honey badgers (Mellivora capensis) and rock pythons (Python sebae). Kittens were found in stomach contents of black-backed jackals in the immediate study area (A. Neils, personal

102

observation). Kitten survival in hunted mountain lion populations was 0.43 (Cooley et al. 2009) to 0.59 (+/-0.21) (Robinson et al. 2008). Decreased kitten survival has been reported for felids in areas with high male turnover when young males enter the population and kill unrelated kittens to bring females into estrus (Sweanor et al. 2000; Cooley et al. 2009; Loveridge et al. 2010; Barthold et al 2016).

4.4. Long-term ecological consequences

Although absolute numbers of caracals on southern Namibian farmlands appears to remain steady, ecological and evolutionary concerns are raised. Chronic low adult survival could cause shifts in reproductive strategy and genetic/ evolutionary trajectories. Individuals within populations can respond to high mortality rates by altering their reproductive strategies, either by increased litter sizes (Knowlton 1972; Sacks 2005; Melero et al. 2015) or reproduction at a younger age (Benton et al. 1995; Ericsson et al., 2001). Caracal litter sizes are reported to range from 2-5 kittens (Smithers 1983; Stuart and Stuart 1985; Weisbein & Mendelssohn 1990). In this study litter sizes averaged 2.2 (+1.09) from the second year onward, thus caracals do not appear to have the innate capacity to increase litter sizes in response to high mortality. Cooley et al. (2009) found that vital rates for puma did not compensate for high hunting mortality. Some mammalian mesocarnivores display shifts in reproductive activity rather than simply increase fecundity

(Knowlton 1972; Sacks 2005; Melero et al. 2015) and some heavily hunted ungulates reproduce and senesce earlier than individuals in populations without low adult survival (Benton et al. 1995;

Ericsson et al., 2001). Evolutionary consequences of reproductive shifts are unknown.

103

Reproduction of some normally iteroparous mammals that experience high mortality is best described as functional semelparity (Goldstein et al. 2017). Since mortality rates are so high, southern Namibian caracals may fit this definition because of the likelihood that only one reproductive effort is made before death at a young age. Many primiparous mothers however produce smaller litters, smaller young, and are less successful at raising young compared with older, more experienced females that often produce the highest quality young (Forslund & Pärt

1995; Smith 2009; Borowik et al. 2016). In this study the highest average litter size was achieved in the 4-5th year followed by the 3-4th year. However, female caracals have only a 16% chance to survive their 3-4th year, and only a 6% chance to survive their 4-5th year, thus caracals rarely have the chance to mature into experienced parents. Poor adult survival inherently constrains the capacity for the population to increase since caracals appear unable to compensate for high mortality with increased reproduction. Reproductive potential of caracals’ iteroparous life history strategy is not achieved under the influence of anthropogenic mortality.

4.5. Conservation implications

Conservation and management of species experiencing heavy anthropogenic impacts can greatly benefit from detailed demographic information. Managers who seek to mitigate anthropogenic impacts must first understand demographic impacts of animal removal. Long-term datasets are needed to gain better insight into caracal population dynamics, survival and reproduction. Southern Namibia could represent a broad population sink; the landscape is saturated with livestock farms whose managers consistently destroy caracals. Identification of sources of immigration is needed since immigration is the likely mechanism sustaining local 104

caracal populations. Outside of farmed areas, natural protected areas may be insufficient sources of individuals for heavily hunted caracal populations as these areas are often saturated with larger predators such as leopard, cheetah and hyenas, which may result in lowered caracal densities (A.

Neils, personal observation). Populations of small felids are low in most protected reserves with higher densities of larger heterospecifics (Caro 1987; Stewart & Wilson 1988). Collaring yearling caracals in protected areas may reveal more clues in regard to the nature of caracal dispersal into neighboring farmed areas.

Southern Namibian caracals are at a crossroads and their future is uncertain. To prevent caracals from joining the ranks of cheetahs, leopard, and lions in southern Namibia, reclassification of caracals from ‘vermin’ to a managed game species like leopard and cheetah may permit populations to rebound with the relaxation of persecution pressure. However, caracal persecution is difficult to curb when the goal of many farmers is to ultimately achieve eradication. Under current Namibian legislation, landowners act within their legal rights to achieve this end. Farmers pressure each other to kill predators, even if the threat is merely percieved (A. Neils, personal observation).

Unfortunately, certain economic industries are often in direct competition (i.e. livestock production versus conservation/ ecotourism), causing conflicts that threaten the integrity of ecological resources. Carnivore populations are prone to signiﬁcant declines as a result of HWC since they require space and resources compromised by increasing human dominance on landscapes, and removal efforts are often the first strategy employed (Zimmerman et al. 2010).

However, predator tolerance can be improved by education and outreach, effective conflict mitigation measures, and promotion of ecotourism as an alternative or supplementary industry 105

(Homern et al. 2007). First and foremost, enhanced scientific understanding of the relationship between human activity and predator ecology is required, as conﬂict resolution requires customized solutions and outreach based on best available ecological data for the species involved as well as the social psychology and economy of the people affected (Ajzen and Fishbein 1980).

Although predator control efforts used by livestock producers have had success in terms of carnivore removal, efforts often fail to prevent industry losses (Berger 2006). Therefore, HWC management must include outreach efforts that influence perceptions of risk (Riley and Decker

2000) and subsequently reduce retaliatory and preventative carnivore persecution. Effective conservation cannot occur without addressing the ecological, economical, and sociological aspects of conflict.

106

LITERATURE CITED

Ajzen, I., Fishbein, M., 1980. Understanding attitudes and predicting behaviour. Prentice Hall, Inc. New Jersey, USA.

Avenant, N.L, Nel, J.A.J., 1998. Home-range use, activity, and density of caracal in relation to prey density. African Journal of Ecology 36, 347-359.

Barthold, J.A., Loveridge, A.J., Macdonald, D.W., Packer, C., Colchero, F., 2016. Bayesian estimates of male and female African lion mortality for future use in population management. Journal of Applied Ecology 53, 295-304.

Beier, P., 1991. Cougar attacks on humans in the United States and Canada. Wildlife Society Bulletin 19, 403-412.

Berger, K.M., 2006. Carnivore-livestock conflicts: effects of subsidized predator control and economic correlates on the sheep industry. Conservation Biology 20, 751–761.

Bernard, R.T.F., Stuart, C.T., 1987. Reproduction of the caracal Felis caracal from the Cape Province of South Africa. South African Journal of Zoology 22, 177-182.

Borowik, T., Wawrzyniak, P., Jędrzejewska, B., 2016. Red deer (Cervus elaphus) fertility and survival of young in a low-density population subject to predation and hunting. Journal of Mammalogy 97, 1671-1681.

Bothma, J.D.P., Le Riche, E.A.N., 1984. Aspects of the ecology and the behaviour of the leopard Panthera pardus in the Kalahari desert. Koedoe 27, 259-279.

Caro, T.M., 1987. Cheetah’s mothers’ vigilance looking out for prey or for predators. Behavioral Ecology Sociobiology 20, 351-362.

Caughley, G., 1966. Mortality patterns in mammals. Ecology 47, 906-918.

Caughley, G., 1977. Analysis of vertebrate populations. John Wiley & Sons, New York.

Clutton-Brock, T.H., Iason, G.R., 1986. Sex ratio variation in mammals. The Quarterly Review of Biology 61, 339-374.

Cockburn, A., Legge, S., Double, M.C., 2002. Sex ratios in birds and mammals: can the hypotheses be disentangled? In: Sex Ratios: Concepts and Research Methods. Edited by Hardy, I.C.W. Cambridge University Press, United Kingdom, pp. 266-286.

107

Cooley, H.S., Wielgus, R.B., Koehler, G.M., Robinson, H.S., Maletzke, B.T., 2009. Does hunting regulate cougar populations? A test of the compensatory mortality hypothesis. Ecology 90, 2913-2921.

Creel, S., 1998. Social organization and effective population size in carnivores. Behavioral Ecology and Conservation Biology, 246-265.

Delibes, M., Ferreras, P., Gaona, P., 2001. Attractive sinks, or how individual behavioural decisions determine source–sink dynamics. Ecology Letters 4, 401-403.

Ericsson, G., Wallin, K., Ball, J.P., Broberg, M., 2001. Age-related reproductive effort and senescence in free-ranging moose, Alces alces. Ecology 82, 1613-1620.

Forslund, P., Pärt, T., 1995. Age and reproduction in birds—hypotheses and tests. Trends in Ecology & Evolution 10, 374-378.

Franke, L.G., Woodroffe, R., 2001. Behaviour of carnivores in exploited and controlled populations. In: Carnivore conservation. Edited by Gittleman, J.L., Funk, S.M., Macdonald, D.W., Wayne, R.K. Cambridge University Press, United Kingdom, pp. 419– 442.

Goldstein, E.A., Merrick, M.J., Kaprowski, J.L., 2017. Functional semelparity drives population dynamics and endangers a peripheral population. Biological Conservation 205, 52-59.

Gordon, I.J., Hester, A.J., Festa-Bianchet, M., 2004. Review: the management of wild large herbivores to meet economic, conservation and environmental objectives. Journal of Applied Ecology 4, 1021-1031.

Holmern, T., Nyahongo, J., Røskaft, E., 2007. Livestock Loss Caused by Predators Outside Serengeti National Park, Tanzania. Biological Conservation 135, 518–526.

Kamler, J.F., Gipson, P.S., Snyder, T.R., 2000. Dispersal characteristics of young bobcats from northeastern Kansas. The Southwestern Naturalist 45, 543-546.

Kok, O.B., Nel, J.A.J., 2004. Convergence and divergence in prey of sympatric canids and felids: opportunism or phylogenetic constraint? Biological Journal of the Linnean Society 83, 527-538.

Knowlton, F.F., 1972. Preliminary interpretations of coyote population mechanics with some management implications. The Journal of Wildlife Management 36, 369-382.

108

Lambert, C.M., Wielgus, R.B., Robinson, H.S., Katnik, D.D., Cruickshank, H.S., Clarke, R., Almack, J., 2006. Cougar population dynamics and viability in the Pacific Northwest. The Journal of Wildlife Management 70, 246-254.

Lindzey, F.G., Van Sickle, W.D., Ackerman, B.B., Barnhurst, D., Hemker, T.P., Laing, S.P., 1994. Cougar population dynamics in southern Utah. The Journal of Wildlife Management 58, 619-624.

Loveridge, A.J., Wang, S.W., Frank, L.G., Seidensticker, J., 2010. People and wild felids: conservation of cats and management of conflicts. Biology and conservation of wild felids, 161.

Marker, L., Dickman, M., 2005. Notes on the spatial ecology of caracals (Felis caracal) with particular reference to Namibian farmlands. African Journal of Ecology 43, 73-76.

Mendelssohn, H., 1989. Felids in Israel. Cat News 10, 2-4.

Moolman, L.C., 1986. Aspects of the ecology and behavior of the caracal Felis caracal Schreber 1776 in the Mountain Zebra National Park and on the surrounding farms. Master’s thesis, University of Pretoria, Pretoria.

Mosby, H.S., 1969. The influence of hunting on the population dynamics of a woodlot gray squirrel population. The Journal of Wildlife Management 33, 59-73.

Mowat, G., Boutin, S., Slough, B.G., 1996. Using placental scar counts to estimate litter size and pregnancy rate in lynx. The Journal of Wildlife Management 60, 430-440.

Mukherjee, S., Goyal, S.P., Johnsingh, A.J.T., Leite-Pitman, M.R.P., 2004. The importance of rodents in the diet of jungle cat (Felis chaus), caracal (Caracal caracal), and golden jackal (Canis aureus) in Sariska Tiger Reserve, Rajasthan, India. Journal of Zoology London 262, 405-411.

Norton, P.M., Lawson, A.B., 1985. Radio tracking of leopards and caracals in the Stellenbosch area, Cape Province. South African Journal of Wildlife Research 15, 17-24.

Nowell, K., Jackson, P., 1996. Wild cats: status survey and conservation action plan. Gland: IUCN, 382.

Peto, R., Peto, J., 1972. Asymptotically efficient rank invariant test procedures. Journal of the Royal Statistical Society Series A, 185-207.

Pringle, J.A., Pringle, V.L., 1979. Observations on the lynx Felis caracal in the Bedford district. South Africa Journal of Zoology 14, 1-4. 109

Pulliam, R.H., 1988. Sources, sinks, and population regulation. The American Naturalist 132, 652-661.

Quan, J., Barton, D., Conroy, C., 1994. A preliminary assessment of the economic impact of desertiﬁcation in Namibia. Directorate of Environmental Affairs, Windhoek. Research Discussion Paper 3, 1–148.

Reid, H., Sahlén, L., Stage, J., MacGregor, J., 2008. Climate change impacts on Namibia's natural resources and economy. Climate Policy 8, 452-466.

Riley, S.J.A., Decker, D.J., 2000. Risk Perception as a Factor in Wildlife Stakeholder Acceptance Capacity for Cougars in Montana. Human Dimensions of Wildlife 5, 50–62.

Robinson, H.S., Wielgus, R.B., Cooley, H.S., Cooley, S.W., 2008. Sink populations in carnivore management: cougar demography and immigration in a hunted population. Ecological Applications 18, 1028-1037.

Rolley, R.E., 1985. Dynamics of a harvested bobcat population in Oklahoma. The Journal of Wildlife Management 49, 283-292.

Ross, P.I., Jalkotzy, M.G., 1992. Characteristics of a hunted population of cougars in southwestern Alberta. The Journal of Wildlife Management 56, 417-426.

Sacks, B.N., 2005. Reproduction and body condition of California coyotes (Canis latrans). Journal of Mammalogy 86, 1036-1041.

Sibly, R.M., Collett, D., Promislow, D.E.L., Peacock, D.J., Harvey, P.H., 1997. Mortality rates of mammals. Journal of Zoology 243, 1-12.

Singh, R., Qureshi, Q., Sankar, K., Krausman, P., Goyal, S.P., 2014. Population and habitat characteristics of caracal in semi-arid landscape. Indian Journal of Arid Environments 103, 92-95.

Skinner, J.D., 1979. Feeding behavior in caracal Felis caracal. Journal of Zoology London 189, 523-525.

Slough, B.G., Mowat, G., 1996. Lynx population dynamics in an untrapped refugium. The Journal of Wildlife Management 60, 946-961.

Smith, H.J., 2009. Parenting for primates. Harvard University Press.

110

Smithers, R.H.N., 1983. The Mammals of the southern African subregion. Pretoria: University of Pretoria.

Strohbach, B.J., 2001. Vegetation survey of Namibia Namibia wissensschaftliche gesellschaft. Namibia Scientific Society Journal 49, 93-124.

Stuart, C.T., 1981. Notes on the mammalian carnivores of the Cape Province, South Africa. Bontebok 1, 1-58.

Stuart, C.T., 1984. The distribution and status of Felis caracal Schreber, 1776. Saugetierkundliche Mitteilungen 31, 197- 203.

Stuart, C.T., Stuart, T.L., 1985. Age determination and development of foetal and juvenile Felis caracal Schreber 1776. Saugetierkundliche Mitteilungen 2, 217-229.

Stuart, C.T., 1986. The incidence of surplus killing by Panthera pardus and Felis caracal in Cape Province, South Africa. Mammalia 50, 556-558.

Stuart, C.T., Hickman, G.C., 1991. Prey of caracal, Felis caracal, in two areas of Cape Province, South Africa. Bontebok 1, 1-58.

Stuart, C.T., Wilson, V.J., 1988. The cats of southern Africa. Zimbabwe: Chipangali Wildlife Trust.

Stuart, C., Stuart, T., 2013. Caracal Caracal caracal: Mammals of Africa 5, 174-179.

Sweanor, L. L., Logan, K.A., Hornocker, M.G., 2000. Cougar dispersal patterns, metapopulation dynamics, and conservation. Conservation Biology 14, 798–808.

Sunquist, M., Sunquist, F., 2002. Wild cats of the world. The University of Chicago Press, Chicago.

Thomas, C.D., Kunin, W.E., 1999. The spatial structure of populations. Journal of Animal Ecology 68, 647-657.

Trainer, C.E., Matson, G., 1988. Age determination in cougar from cementum annuli counts of tooth sections. Proceedings of the Mountain Lion Workshop, 3, 71.

Van Heezik, Y.M., Seddon, P. J., 1998. Range size and habitat use of an adult male caracal in Northern Saudi Arabia. Journal of Arid Environments 40, 109-112.

Visser, J., 1976. Status and conservation of the smaller cats of southern Africa. In: The World’s Cats, vol. 3 ed. R.L.Eaton. Seattle: Carnivore Research Institute, 60-66. 111

Weisbein, Y., Mendelssohn, H., 1990. The biology and ecology of the caracal (Felis caracal) in the northern Aravah Valley of Isreal. Cat News 12, 20-22.

Wielgus, R.B., Morrison, D.E., Cooley, H.S., Maletzke, B., 2013. Effects of male trophy hunting on female carnivore population growth and persistence. Biological Conservation 167, 69- 75.

White, G.C., 2000. Population viability analysis: Data requirements and essential analyses. In: Research techniques in animal ecology: Controversies and consequences. Edited by Boitani, L., Fullers, T.K. Cambridge University Press, New York, 288-331.

Woodroffe R., Ginsberg, J.R., 2005. King of beasts? Evidence for guild redundancy among large mammalian carnivores. In: Large Carnivores and the Conservation of Biodiversity. Edited by Ray, J.C., Redford, K.H., Steneck, R.S., Berger, J. Island Press, 154-175.

Zimmermann, A., Baker, N., Inskip, C., Linnell, J.D., Marchini, S., Odden, J., Rasmussen G., Treves, A. 2010. Contemporary views of human–carnivore conflicts on wild rangelands. In Wild rangelands: Conserving wildlife while maintaining livestock in semi-arid ecosystems, 129-151.

112

TABLES

Table 1- Numbers of yearling/ sub-adult (Y/ SA) to adult (A) caracals (Caracal caracal) on southern Namibian farmlands monitored from 2010-2016.

Collared Necropsied % Young animals % Young animals Y/ SA A Y/ SA A Total population M 15 5 66 29 70.4 39.1 F 5 9 45 33 54.3 24.1 Total 20 14 111 62 63.2

113

Table 2- Life table statistics calculated for all caracals (Caracal caracal) of known age from southern Namibian farmlands collected from 2010-2016.

Age class Number Number Death Survival Midpoint Expected # age classes (yrs) Alive Survivorship Dying Rate Rate Survivorship remaining x nx lx dx qx sx Lx Tx 0 208 1.000 33 0.159 0.841 0.921 2.462 1 175 0.841 99 0.566 0.434 0.603 1.541 2 76 0.365 41 0.539 0.461 0.267 0.938 3 35 0.168 21 0.600 0.400 0.118 0.671 4 14 0.067 10 0.714 0.286 0.043 0.553 5+ 4 0.019 4 1.000 0.000 0.019 0.510

114

Table 3- Life table statistics calculated for all male caracals (Caracal caracal) of known age from southern Namibian farmlands collected from 2010-2016.

Age class Number Number Death Survival Midpoint Expected # age classes (yrs) Alive Survivorship Dying Rate Rate Survivorship remaining x nx Lx dx qx sx Lx Tx

0 115 1.000 18 0.157 0.843 0.922 2.462 1 97 0.843 62 0.639 0.361 0.574 1.540 2 35 0.304 15 0.429 0.571 0.239 0.966 3 20 0.174 11 0.550 0.450 0.126 0.727 4 9 0.078 7 0.778 0.222 0.048 0.601 5+ 2 0.017 2 1.000 0.000 0.017 0.553

115

Table 4- Life table statistics calculated for all female caracals (Caracal caracal) of known age from southern Namibian farmlands collected from 2010-2016.

Age Number Survivorship Number Death Survival Midpoint Expected # Avg. # class Alive Dying Rate Rate Survivorship age classes female (yrs) remaining young*

x nx lx dx qx sx Lx Tx bx

0 92 1.000 15 0.163 0.837 0.918 2.462 0.000

1 77 0.837 35 0.455 0.545 0.647 1.544 0.286

2 42 0.457 27 0.643 0.357 0.310 0.897 1.042

3 15 0.163 9 0.600 0.400 0.114 0.587 1.346

4 6 0.065 4 0.667 0.333 0.043 0.473 1.584

5+ 2 0.022 2 1.000 0.000 0.022 0.429 1.250

r0 = 1.064

*An even female: male sex ratio of kittens was assumed

116

Table 5- Life table statistics calculated for all female caracals (Caracal caracal) of known age from southern Namibian farmlands collected from 2010-2016.

Age Number Survivorship Number Death Survival Midpoint Expected # Avg. # class Alive Dying Rate Rate Survivorship age classes female (yrs) remaining young*

x nx lx dx qx sx Lx Tx bx

0 92 1.000 15 0.163 0.837 0.918 2.462 0

1 77 0.837 35 0.455 0.545 0.647 1.544 0.239

2 42 0.457 27 0.643 0.357 0.310 0.897 0.872

3 15 0.163 9 0.600 0.400 0.114 0.587 1.127

4 6 0.065 4 0.667 0.333 0.043 0.473 1.325

5+ 2 0.022 2 1.000 0.000 0.022 0.429 1.046

r0 = 0.891

*Female: male sex ratio of kittens based on observed sex ratios of adult and subadults in the population

117

Table 6- Kaplan-Meier survival curve statistics (days) for male, female, yearling/ subadult, and adult radio-collared caracals (Caracal caracal) from three different habitat types (Karoo/

Karasberg, Khalahari grassland; sparse shrubland) on southern Namibian farmlands from 2010-

2016.

Factor Mean SE 95% CI for the mean Median 95% CI for the median Female 169.357 54.45 62.634 to 276.080 49 24.000 to 287.000 Male 66.5 18.737 29.775 to 103.225 23 21.000 to 83.000 Karoo/ Karasberg 92.273 45.627 2.843 to 181.702 49 21.000 to 90.000 Kalahari grassland 108.143 39.574 30.577 to 185.709 26 17.000 to 140.000 Sparse shrubland 130.222 57.61 17.308 to 243.137 42 22.000 to 117.000 Yearling/ sub-adult 44 16.832 11.008 to 76.992 22 9.000 to 33.000 Adult 201.5 49.624 104.237 to 298.763 106 83.000 to 287.000

118

FIGURE LEGENDS

Figure 1- Kaplan-Meier survival curve for male and female radio-collared caracals (Caracal caracal) from southern Namibian farmlands monitored from 2010-2016 (P = 0.0573).

Figure 2- Kaplan-Meier survival curve for yearling/ subadult (Y/ SA) and adult (A) radio-collared caracals (Caracal caracal) from southern Namibian farmlands monitored from 2010-2016 (P =

0.0002).

Figure 3- Kaplan-Meier survival curve for radio-collared caracals (Caracal caracal) from three different habitat types (Karoo/ Karasberg, KK; Khalahari grassland, KG; sparse shrubland, SS) on southern Namibian farmlands monitored from 2010-2016 (P = 0.9615).

119

FIGURES

Figure 1- Kaplan-Meier survival curve for male and female radio-collared caracals (Caracal caracal) from southern Namibian farmlands monitored from 2010-2016 (P = 0.0573).

120

Figure 2- Kaplan-Meier survival curve for yearling/ subadult (Y/ SA) and adult (A) radio-collared caracals (Caracal caracal) from southern Namibian farmlands monitored from 2010-2016 (P =

0.0002).

121

122

APPENDIX C: UNINTENDED CONSEQUENCES: HOW THE ANIMAL

RIGHTS MOVEMENT INADVERTENTLY LED TO THE PERSECUTION

OF NAMIBIAN CARNIVORES

Aletris M. Neils

(In the format of Human Ecology)

123

UNINTENDED CONSEQUENCES: HOW THE ANIMAL RIGHTS MOVEMENT

INADVERTENTLY LED TO THE PERSECUTION OF NAMIBIAN CARNIVORES

Aletris Marie Neils1,2 1Conservation CATalyst, Tucson Arizona 2University of Arizona, Tucson Arizona

ABSTRACT

Namibia has a long history of human-wildlife conflicts and predator persecution. The objective of this research was to use qualitative interviews to enhance understanding of wildlife conflicts and risk perception from a socio-economic context with the goal of identifying management actions to reduce the number of predators killed by people. Respondents indicated that drivers of conflict include drought, resource availability, and market-driven demands for newer breeds of sheep whose husbandry practices leads to increased conflicts with predators. Animal rights campaigns in the 1980s led to a crash in the once profitable market for localized Swakara® pelts from Karakul sheep. The shift to mutton breeds led to increased conflicts with native predators now persecuted in higher numbers than ever. Results suggest farming systems were created where predator eradication is the strategy over livestock management. This scenario demonstrates how well-intended political campaigns can have unintended consequences on ecological and social systems.

KEYWORDS: jackal, caracal, indigenous knowledge, human-carnivore conflict, Namibia, wildlife perceptions

124

INTRODUCTION

Conﬂicts between wildlife and people pose a significant challenge to conservationists

(Woodroffe & Ginsberg 2005). Livestock depredation and subsequent predator control responses by farmers are a growing concern worldwide (Loveridge 2010). Human–wildlife conﬂict (HWC) arises when a species poses a direct and recurring threat to community livelihood. HWC occurs in many different habitats, but rangelands have always been particularly prone to conﬂicts; this is often where carnivores, prey, people and livestock occur together in the same space and are likely to encounter one another (Zimmerman 2010). Much of

Namibia’s agricultural land (44%) is utilized for livestock production (Schneider 1994).

Namibian farmlands support a combination of domestic livestock (cattle, goats, sheep, and poultry) and game but are also home to the majority of Namibia’s wildlife (Marker et al. 2003).

Magnitude of HWC can be real or perceived, but often results in persecution efforts of predators (Zimmerman et al. 2010). Persecution is defined as the persistent killing, chasing or other harassment of a species with the goal of eradication, and includes both retaliatory and preventative killings (Zimmerman et al. 2010). Persecution is common among commercial ranchers that lose livestock to predation (Zimmerman et al. 2010). Livestock owners who experience high depredation are more approving of lethal measures, and often HWC causes direct opposition to conservation efforts (Holmern et al. 2007).

Populations of large carnivores are prone to signiﬁcant declines as a result of HWC since they require space and resources compromised by increasing human dominance on landscapes

(Zimmerman et al. 2010). HWC often occurs where natural prey is depleted and livestock provides an alternative food source (Woodroffe & Ginsberg 1998). In southern Namibia, large 125

carnivores such as lion (Panthera leo), African wild dog (Lycaon pictus), and spotted hyena

(Crocuta crocuta) have already been systematically extirpated due to persecution from farmers following European settlement. Only fragmented remnant populations of leopard (Panthera pardus) and cheetah (Acinonyx jubatus) remain. Mesocarnivores such as caracals (Caracal caracal) and black-backed jackals (Canis mesomelas) still remain but are now considered the greatest predatory threat to small stock such as sheep and goats by local farmers (Stuart &

Hickman 1991; Mukherjee et al. 2004; Stein 2006). Carnivore persecution in southern Africa has occurred on a large scale for decades (Stuart & Hickman 1991; Stuart & Stuart 2013). For example, an average 185 caracals are killed annually on the farms surrounding Zebra National

Park in South Africa (Moolman 1986). From 1931-1952, over 2,200 caracals per year were killed in control operations in the Karoo, South Africa (Stuart 1981). Similarly, farmers in

Namibia reported killing up to 2,800 caracals in 1981 alone (Nowell and Jackson 1996). This pace of persecution may have lasted for decades on a regional scale and likely continues to this day, although no recent data exist. Predator control efforts, however, often fail to prevent declines in the livestock industry (Berger 2006). The goal of this study was to investigate human dimensions of HWC in Namibia to better understand causative social and economic reasons that underlie continued and often rampant carnivore persecution.

METHODS

2.1. Study site

This research was conducted on privately owned commercial livestock farmlands in

Namibia, from the veterinary cordon fence (VCF) south to the Orange River. The region is arid, 126

with the southern portions being extremely arid, with highly fluctuating and unpredictable rainfall (Strohbach 2011). Large tracts of grasslands are relatively scarce and insufficient to support large scale cattle farming; therefore, commercial farmers rely on small stock, primarily sheep and goats (Quan et al. 2004). To a lesser extent, wild game such as kudu (Tragelaphus strepsiceros), oryx (Oryx gazelle) and springbok (Antidorcas marsupialis) are used by landowners for subsistence and commercial hunting. This study focused on farms south of the

VCF, because areas north of the VCF cannot sell livestock internationally due to concerns of

Rinderpest and foot-and-mouth disease outbreaks (Miescher 2012). Purposeful sampling efforts were made to select farms south of Windhoek, since these areas typically have the lowest amount of rainfall and primarily farm with small stock.

2.2. Subjects

Qualitative interviews were conducted with commercial Namibian farmers, primarily recruited from farmers’ associations meetings and a personal network developed since 2002. All farmers were interviewed personally by the author. After initial participants were identified, snowball and purposeful sampling were used to find farms to visit and farmers to interview.

Farmers were primarily of Afrikaaner or German descent.

2.3 Interviews

Qualitative data concerning HWC and broader landscape issues of the study area were collected via semi-structured interviews. At least one interview with each farmer included a list of questions and topics discussed. Additional interviews with farmers were mostly unstructured, some taking place in an unplanned and informal ad hoc manner (primarily outside on the farm, in

127

corrals, or after agricultural associate meetings). Informed consent was obtained from participants before each interview occurred (University of Arizona Human Subjects Protection

Permit #1510165011).

Interviews were conducted until theoretical saturation was reached (Wutich and Gravlee

2010). In addition to collecting socio-demographic data, interviews explored respondents’ experience with carnivores, their perceptions of wildlife, and income from farming and natural resources. Interviews had four main focuses:

1. Identification of carnivores & other wildlife on the farmer’s property,

2. Livestock husbandry and losses due to predators

3. Gathering opinions of what factors contribute to their conflict with predators

4. Methods used and numbers of predators killed by farmers.

Interviews also included a Participatory Risk Mapping (PRM) section comprising open questions about the problems the respondent and their household members face while farming and coexisting with predators. To avoid leading the respondents, the PRM section of the questionnaire preceded all questions concerned specifically with carnivores and human-wildlife conflict (Inskip et al. 2013).

Additional subjects were farmers who contacted the author when predators were killed on their property or who reported livestock losses due to suspected predators. The author would either travel to their farm for forensic investigations, or carcasses were frozen and stored by farmers to be collected by the author at a later date. During these investigations, farmers were interviewed in an ad hoc, qualitative manner about livestock husbandry, predators (losses and removal), determination of losses, and threats farmers face. The author also accompanied farmer 128

and/ or farm workers into the veldt to see killsites, the farm, predator control methods, and livestock.

2.4 Analysis

All interviews and conversations were transcribed onto MAXQDA Analytics Pro

(VERBI Software GmbH, Berlin, Germany) for further qualitative analysis. Each entry was read and data were then coded into themes using a grounded-theory approach. Quotes used in the results section were selected for their typical representation of a particular theme that emerged from the data (Auerbach and Silverstein 2003).

RESULTS

A total of 561 qualitative interviews were conducted with 367 Namibian farmers. Of these, 482 interviews were conducted with 309 farmers (13 female, 296 male) that contacted the author with a carnivore conflict scenario and/or carnivore carcass. To representatively sample all farmers, not only those experiencing conflicts, an additional 79 in-depth, qualitative, semi- structured interviews were conducted with 58 farmers (7 female, 51 male). Semi-structured interviews lasted between 19 minutes and 178 minutes, averaging approximately 90 minutes.

Farmer age ranged from 33-87 years old, with a mean of 57. Nearly half of the farmers interviewed were of German decent, 40% of Afrikaans decent, 10% identified as having a mixed background.

3.1. Farm characteristics-

Farms averaged approximately 11,000 ha (ranged <3000 to > 30000). Some farmers

(19%) also rent/hire adjacent farms (or portions of farms) to hold more livestock. Farmers 129

reported having an average of approximatly1600 head of livestock (range 130- >4700). Farmers reported making between approximately 40% and 100% of their income from livestock farming, however >70% of farmers reported that they also made some income from game meat, mostly through hunting springbok to produce ‘biltong’ (strips of dried meat) in the winter. Some farmers also had guest farms, shops, and/or had weekday jobs in cities. All farmers reported they wanted to continue farming. However 21% were interested in pursuing or expanding for ecotourism and/or trophy hunting. Eighty-nine percent plan to keep the farm in the family by bequething to their children or grandchildren. Seven percent plan to sell their farm in the future.

3.2. Livestock and predation-

Farmers interviewed can be divided into pelt farmers (those raising Karakul sheep) or mutton farmers (those raising Dorper, Von Roy, or hybrid varieties such as ‘meat masters’). The majority of mutton farmers (64%) acknowledged that they (or their family) used to farm with

Karakul sheep until demand decreased in the 1980s. Most mutton farmers (79%) acknowledged that Karakul sheep had less of an environmental impact and could be better managed against predators. However, only 6% of mutton farmers said would consider reverting to Karakul production.

Mutton farmers reported higher losses to predators, higher amounts of predators killed or removed from their farms, and a higher amount of time and money spent on predator removal than pelt farmers. Most farmers (89%) said that certain livestock breeds were less susceptible to predation. Of these Nguni cattle were the large stock most often recognized as least likely to be taken by predators and as the livestock breed that has the least impact on the environment. Most 130

farmers (84%) reported that small stock was “easier on the veldt” or had less of an environmental impact than cattle.

In terms of livestock losses, 64% said drought was the time of the greatest loss. The top driver of predator persecution actions was linked to drought and/or climate change; the greatest perceived threat farmers face is drought/ lack of rain. Fifty-three farmers (91%) acknowledged this problem is worse than it used to be, and thirty-three farmers (56%) specifically mentioned

“climate change” or “global warming.” All farmers over 70 years of age mentioned significant climate changes within their lifetime and 67% of these farmers expressed concerns about the climate impacting the future viability of raising livestock on their farm and/or in Namibia.

Seventy-two % of Karakul sheep farmers interviewed were under 50 years of age (many under 45), suggesting that the younger generation of farmers is returning to farming with this breed. While a majority (86%) of farmers over 50 years of age were dorper or mutton sheep farmers. More young farmers (45 years or less) stated being receptive/interested in learning strategies for coexisting with predators and mitigating conflicts (other than removal) than older farmers (50 years or older). Interestingly, senior farmers (70+ years) also expressed regret when talking about past predator removals.

One third of farmers said the greatest time of stock loss was during breeding season. A significant number of farmers (18%) specifically mentioned that the greatest livestock losses occur in association with jackals pupping. The top small stock that were reported as the least likely to be taken by predators were the two pelt breeds, Karakul sheep (75%) and Damara sheep

(22%), likely a result of the innate flocking behavior of these breeds and associated husbandry techniques. 131

Overall, black-backed jackals were viewed by farmers as the top problem predator (44%), followed by caracal (21%), cheetah and/or leopard (17%), baboon (9%), people (6%), or other

(3%). Self-reported average number of predators removed on mutton sheep farms (Dorper, etc.) per year was 61 (range 4-376), while the average number of predators removed on Karakul sheep farms was 6 (range 0- 141). Mean number of predators killed was significantly different between flock sheep farmers and mutton sheep farmers (t = -3.06, df = 15, P < 0.008). All

Dorper farmers reported removing at least one predator within the last year compared to 73% of pelt sheep farmers. The top methods used for removing predators on Dorper farms were spotlight shooting and setting gin (steel-jaw leg-hold) traps, whereas Karakul farms typically used dogs for predator deterrence or removal.

When farmers were asked how they managed conflicts with predators, 91% of pelt farmers first mentioned strategies of actively protecting livestock from predators (using shepherds, guarding dogs, corrals). Conversely, 87% of mutton farmers managed conflicts by eradicating predators (traps, poison, shooting, etc.).

There is an apparent paradigm difference determined by sheep variety being raised.

Karakul farmers focus on managing their flocks whereas mutton farmers focus on managing predators to eliminate threats to unmanaged livestock (Figure 1). Almost all Karakul farmers

(>94%) reported using a shepherd to protect the flock, whereas few mutton farmers utilized this technique (<8%). Most Karakul farmers (86%) reported building corrals to protect livestock from predators, compared with relatively few mutton sheep farmers (17%). Over 91% of Dorper sheep farmers utilized either jackal-proof fences, game fences, or electric fences in some part of their farm, compared to 23% of Karakul farmers. Almost one third of Karakul farmers reported 132

they did not have fences and the remainder reported having basic strand stock fences rather than jackal-proof fences. Permeability of these farms were different for wildlife, with mutton farms having barrier fences (game proof, electric, jackal proof, etc.) more than karakul farms (chi- squared= 14.243; P = 0.0002).

DISCUSSION

Understanding human dimensions of HWC and causative factors that lead to widespread indiscriminate persecution is paramount to developing management strategies and allows emphasis on important areas of conflict and to identify possible factors of influence (Holmern et al. 2007).

4.1. Predator perceptions and persecution-

Predator removals were associated with perceived predator problems in interviews, but many indiscriminate removals also occurred. One explanation is that farmers pressure each other to kill predators, even if the threat is merely perceived. Perceived risks often greatly exceed estimates of actual risks (Riley and Decker 2000), and reported magnitude of depredations can be exaggerated. Persecution is economically unjustified as it usually does not ultimately result in reduced conflicts with livestock (Berger 2006). Tolerance of predators is perhaps dictated or influenced by financial well-being with an indirect relationship between margins of profit and persecution efforts.

133

4.2. Effects of culling on carnivore populations

In most cases a combination of factors leads carnivores into depredation and impacted ranchers or pastoralists into retaliatory or preventative persecution. In some rangelands such as those in southern Namibia, predators are persecuted to the point of eradication, presenting both ecological and ethical issues. Carnivores are fundamentally important top-down regulators of herbivore communities and their removal from ecosystems have wide implications for ecosystem structure and function (Treves & Karanth 2003; Woodroffe & Ginsberg 2005; Treves et al. 2006;

Zimmerman 2010).

Effects from culling on carnivore populations include altered population dynamics (Sacks

2005), increased infanticide (Stuart 1981), and decreased genetic diversity due to reduced effective population size. Since carnivores often serve as top-down ecosystem regulators, potential ecosystem effects of carnivore removal include meso-predator relief (overpopulation of mid-sized carnivores), overpopulation of prey species, and increased disease (Woodroffe &

Ginsberg 2005). Namibia has one of the largest remaining populations of free ranging caracals, most of which reside on commercial farms (Marker et al. 2003). Caracals, along with black- backed jackals, are primary targets of persecution efforts in Namibia.

Since many carnivores are territorial, they prevent conspecifics and other predators from entering their territory. Felids including caracals are strongly territorial (Stuart 1981). Increased immigration of young male felids following resident removal has been observed for bobcats

(Lynx rufus), pumas (Puma concolor) and African lions (Rolley 1985; Ross & Jalkotsky 1992;

Slough & Mowat 1996; Sweanor et al. 2000; Kamler et al. 2000; Lambert et al 2006; Cooley et al. 2009; Barthold et al. 2016). Frequent deaths of adult male pumas and African lions are 134

known to disrupt social structure of the population (Sweanor et al. 2000; Barthold et al. 2016).

Increased immigration and recruitment of young animals from adjacent areas results in little or no change in densities, but shifts population structure towards young inexperienced animals

(Robinson et al. 2008). If dominant residents are selecting wild prey, they prevent stock loss.

However, when animals are arbitrarily and indiscriminately removed, as is the case with most methods of predator removal, the territory is rendered vacant and an influx of conspecifics and other predators will quickly fill this void (Robinson et al. 2008). Conspecifics that recolonize a vacant territory are often young and inexperienced hunters. These individuals are actually more likely to take livestock, since they are easier to kill than wild ungulates (Zimmerman 2010).

The southern Namibian caracal population may be experiencing constant influx and rapid turnover of young males. By using non-selective predator culling, many farmers may be increasing their stock losses by creating ecologically unstable carnivore communities.

In order to avoid indiscriminate predator control, stockraisers must first accurately identify sources of stock mortality, and if caused by a predator must be able to undeniably identify the individual culprit. Whether a predator actually killed stock or is merely scavenging must also be discerned (Halbritter et al. 2008). Once an individual predator is identified, it must be tracked and trapped. Such action must be taken immediately and requires prompt knowledge of predation events; often such rapid response of depredation experts is difficult or unfeasible.

The longer the time lag, the more difficult it becomes to identify and subsequently remove the problem animal; this is particularly true for species that have vast home ranges (A. Neils, unpublished data).

135

4.3. Legal ownership of wildlife-

In Namibia, farmers own wildlife that exists on their property (Muir-Leresche and Nelson

2000). In some ways this encourages farmers to be more conservation oriented. For example, if they want to harvest wild ungulates such as kudu, springbok, and oryx, they need to ensure they have sufficient forage and water available on their land, so the animals remain on the property.

On the other hand, farmers are free to do as they choose to wildlife they consider pests, such as predators, without any reporting. To remove predators, farmers use a variety of different methods including poisons, spotlight shooting, hunting dogs, gin traps, remote fence cannons, and live traps. While actively removing predators that are considered to be responsible for the most stock losses, such as caracal and black-backed jackals, many other species are killed including bat-eared fox (Otocyon megalotis), aardwolf (Proteles cristata), aardvark

(Orypteropus afer), porcupine (Hystrix cristata) and warthogs (Phacochoerus africanus) (A.

Neils, unpublished data). Aardvarks, porcupines, warthogs and bat-eared fox are also intentionally killed because they dig holes under fences which other animals can access (A.

Neils, personal observation). Use of distributed poisons and poisoned bait piles may kill a host of species including hawks, eagles, owls, foxes, genets (Genetta genetta), black-footed cats

(Felis nigripes), African wildcats (Felis sylvestris lybica) and mongoose (Herpestidae) (Ogada

2014). Farmers utilizing indiscriminate methods such as spotlight shooting may inadvertently kill anything with reflective eyes, such as small ungulates like steenbok (Raphicerus campestris), large ungulate calves (primarily kudu), and occasionally even the sheep they are trying to protect

(A. Neils, unpublished data).

136

4.4. Drought and climate change-

The top driver of predator persecution actions was linked to drought and/or climate change; the greatest perceived threat farmers face is drought/ lack of rain. Fifty-three farmers

(91%) acknowledged this problem is worse than it used to be, and thirty-three farmers (56%) specifically mentioned “climate change” or “global warming.” Namibian habitats are known to be shifting in response to climatic fluctuations, and drought is an increasing problem throughout much of the country (Strohbach 2001)

Carnivore populations are particularly prone to signiﬁcant declines as a result of HWC since they require spaces and resources compromised by increasing human dominance on landscapes (Zimmerman et al. 2010). Drought conditions exacerbate such resource competition, and can cause declines in native prey. HWC often occurs where natural prey is depleted and livestock provides an alternative food source (Woodroffe & Ginsberg 1998). An individual predator’s health and condition also correlates with livestock depredation (Rabinowitz 1986;

Linnell et al.1999; Miquelle et al.1999; Wydeven et al. 2004); for many individuals, physical condition is sub-optimal during prolonged droughts.

4.5. Livestock breeds-

Livestock management practices often affect frequency and severity of HWC

(Hoogesteijn 2003; Marker et al. 2003; Ogada et al. 2003). One of the primary drivers of carnivore persecution was the type of livestock used on farms. Across Namibia, different breeds of small stock are used commercially, each with varying needs, behaviors, husbandry requirements, and human utilization. Two general categories of small commercial stock were

137

identified, those raised for wool or pelts (pelt sheep) and those raised for meat (mutton sheep).

Due to inherent differences due to biology, behavior, and management strategies, some breeds and their associated husbandry are more prone to HWC.

Wool sheep- Karakul sheep, also known as “Persian lambs” or “Broadtail” are native to

Central Asia and one of the oldest breeds of domesticated sheep (Oklahoma State University

2015). Karakul are hardy and well-adapted to desert environs, storing fat in their distinctively large tail for nourishment in lean times. The wool of the adult Karakul, a very strong fiber, was felted or spun into fabric for garments, footwear, carpets, and yurts, among other uses (Shoeman

1998). Karakul were traditionally the hallmark breed raised for market in Namibia since 1907

(Bravenboer 2007). Although known as the "fur sheep", Karakul are raised for textured, silky pelts taken from lambs harvested at 1-2 days of age. The pelt of the lamb is a lustrous coat of intricately patterned curls; in Namibia Karakul pelts are known as Swakara®1, a modified abbreviation for South West African Karakul Association (Shoeman 1998). Karakul breeding was an important source of income and historically was the most profitable sheep industry and in

Namibia accounts for annual export earnings of 4.5 to 5.5 million Euros (Bravenboer 2007). The industry employs close to 20,000 people in primary and satellite industries. Given Namibia's

1 Swakara® is a registered brand name originating from the arid regions of Namibia. In areas so dry that conventional farming is impossible, the Swakara® industry provides communities with essential income. Production is in harmony with these fragile ecosystems and Swakara® therefore represents a sustainable resource. Animal welfare is strictly regulated by the Swakara® Board of Namibia’s Code of Practice, a set of internationally- recognised ethical standards that are enforced by law. Swakara® is certified Origin Assured ™ by the International Fur Trade Federation, a further guarantee that these important standards are met.

138

social structure, most people working in the industry are families whose sole source of income are Karakul breeding (Shoeman 1998).

On communal lands, small stock is typically represented by Damara sheep, originally from Eastern Asia and Egypt but localized varieties named after Damaraland in Namibia (Du

Toit 2008). Damara sheep have Karakul ancestry and are a dual-purpose breed; the skins of adults are tanned and the meat is consumed. They are found in northwestern Namibia

(Kaokoland) and southern Angola where they were herded by local inhabitants (Himba and

Tjimba) and are relatively free from external influences (Du Toit 2008). The Damara is also a fat-tailed sheep that grows short, coarse hair. They have a diverse diet, feeding on grass, bushes and shrubs as a browser; up to 64% of the diet of the Damara sheep can consist of browsing material (DAD-IS 2017). Damara sheep can survive in harsh environments under poor nutritional conditions and can reproduce where water and grazing is fairly restricted. They have high resistance to most sheep diseases and tolerance against internal parasites (Oklahoma State

University 2015). The reproductive ability of Damara sheep is exceptional. Ewes produce enough milk even to raise twin lambs which will occur in 5 to 10% of the births (Du Toit 2008).

They care well for their young and are known to confront predators when attacked (A. Neils, unpublished data). These features make the Damara breed a good fit for the climate and landscape of Namibia, but carcass weights are small rendering the breed a poor fit for commercial meat production.

Mutton sheep- Dorper sheep, with their characteristic black head and an all-white body, were developed in the 1930s in South Africa as a large meat sheep with a fast growth rate and high reproductive rate (Lategan 2004). Dorper lambs can grow up to 91g daily and can reach a 139

weight of over 36kg in just 4 months (Oklahoma State University 2015). They are highly fertile and can reproduce year-round (Oklahoma State University 2015). Dorper sheep require close management in order to synchronize herd-wide lambing, but can lamb every 8 months if conditions are favorable (Lategan 2004). Dorper sheep are relatively adaptable in terms of climate, grazing conditions, and available forage types (Du Toit 2008). Dorper skin is also a commodity for market and is used to make products requiring thin and strong leather, such as gloves. Skin represents up to 20% of Dorper carcass value (Oklahoma State University 2015).

Dorper sheep are one of the top breeds of commercial mutton in southern Africa and are increasingly common in many countries around the world including the U.S (Lategan 2004).

Less common mutton sheep used in Namibia are Black-headed Persians and Van Rooys

(Du Toit 2008). Black-headed Persians, also known as Swartkoppersie, originated in Somalia is one of the parent breeds of the Dorper (Du Toit 2008). It a fat-rumped breeds and both sexes are polled2. The Van Rooy, also known as the White Persian, is an all-white sheep native to South

Africa first developed in 1906 by J. C. van Rooy, a South African Senator and farmer in the

Bethulie district. The Van Rooy is a cross between indigenous Ronderib Afrikaner sheep,

Blackhead Persian, and Rambouillets (Oklahoma State University 2015); it is a fat-tailed sheep very well-suited to arid climates. Van Rooys are generally kept for meat production but are also used for wool (Du Toit 2008). Like Black-headed Persians, Van Rooys are also polled.

2 Polled livestock are livestock without horns in species which are normally horned. The term refers both to breeds or strains which are naturally polled through selective breeding and also to naturally horned animals which have been dehorned. Natural polling occurs in cattle, yaks, water buffalo and goats, and in these it affects both sexes equally; however, in sheep, both sexes may be horned, both polled, or only the females polled. 140

4.6. The Anti-Fur Movement’s impact to Namibian farmers-

Market profitability and public demand for livestock products generally determine which breeds are invested in and utilized by southern Namibian farmers for livelihoods. Product information from outside sources has a significant and measurable effect on consumer attitudes and purchase actions (Lee 2014). Until the 1980s, a majority of Namibian small stock on commercial farms was Karakul primarily raised for pelts, although the economy was also supplemented by a handful of breeds used for meat. Anti-fur campaigns became part of popular culture during the 1980s-1990s beginning with the anti-sealing campaign in the 1970s and expanding to a more general campaign focused on all animals used to make fur garments

(Emberley 1997). According to the Food and Agricultural Organization of the United Nations

(2017) the minimum population of karakul sheep in Namibia in 1980 was nearly 12 billion.

However, within a few years the market crashed and farmers raised 10.6 billion fewer karakul by

1985 (Table 2).

Animal rights groups began campaigns to bring awareness of the anti-fur movement to the general public in hopes of discouraging consumers from purchasing fur products. Lynx, a

British animal rights organization, used mass media to bring the anti-fur movement to the center of pop culture. Their print ads and videos had a huge impact on the general population, targeting middle- to upper- class Caucasian women as the main consumers of fur products (Emberley

1997). Ads focused on shock value and were designed to make wearing fur be viewed as morally wrong instead of a status symbol. With slogans such as “It takes up to 40 dumb animals to make a fur coat; but only one to wear it” (Lynx anti-fur poster 1984), and posters showing

141

animals caught in steel traps, the public began to see fur in a new way (Emberley 1997). Many celebrities became involved in these anti-fur politics and continue to be used as icons for anti-fur campaigns. Famous fashion icons such as Brigitte Bardot were able to exert a strong influence on the public purchasing fur as a fashion accessory. Bardot, other celebrities, and many other individuals dedicated to exposing the fur trade seemed to change the consumer’s opinion on wearing furs. In Britain, fur sales fell 75% between 1985 and 1990 (Emberley 1997).

People for the Ethical Treatment of Animals (PETA) aggressively launched its anti-fur efforts/ campaigns in the 1980s. These included ads of naked models and celebrities with the slogan “I’d rather go naked than wear fur.” PETA focused heavily on press coverage and their media campaigns became very well known. Pictures of celebrities exposing their bodies in protest of wearing fur attracted the attention of not only anti-fur enthusiasts but many pop-culture followers. Greenpeace International also launched an anti-fur campaign in 1984 (Emberley

1997). Their aggressive campaign included famous fashion photographers and numerous celebrities. However, when controversy over this campaign’s effect on indigenous trappers of

Northern Canada came to light, Greenpeace cancelled this campaign (Harter 2004). Such unintended consequences of far-reaching social campaigns were also felt in Namibia. After the demand of fur decreased, the value of Karakul sheep decreased and the impact on Namibian sheep farmers was tremendous.

Until the aforementioned anti-fur campaigns in the 1980s, Karakul sheep were the primary small stock for commercial farms in Namibia with pelts from Namibia sold to couture fashion markets in Europe. However, methods used for harvesting pelts made the industry very susceptible to the scrutiny and impact of the anti-fur movement. To maximize the value of 142

Karakul pelts, lambs are harvested between 1-3 days old while hair is short, textured and still very glossy. Under a changing social perspective, these methods were deemed barbaric and the market in Europe quickly crashed. While adult Karakul still had a value for mutton, the carcass weight is significantly smaller than that of meat-specific breeds. Native Damara sheep, although very versatile, also did not produce a carcass weight that could support profit for competitive commercial production of meat. In response, Namibian commercial farmers were forced to switch to raising meat specific breeds for market, and this shift had a tremendous impact on native wildlife.

4.7. Conservation implications of market shift-

Several aspects of the behavior and husbandry techniques between Karakul and mutton breeds led to clash with native predators and other wildlife after this market shift. Vital management distinctions exist between wool sheep and mutton sheep industries with direct consequences for native predators. Ironically, the animal rights and anti-fur movements inadvertently caused an enormous increase in the number of native carnivores killed as backlash for HWC.

Flock behavior and management- Karakul farmers focus on managing their flocks whereas mutton farmers focus on managing predators to eliminate threats to unmanaged livestock. Almost all Karakul farmers (96%) reported using a shepherd to protect the flock, while few mutton farmers utilized this technique (8%), as the high value of Karakul pelts made hiring shepherds economically viable. Most Karakul farmers (86%) reported building corrals to protect livestock from predators, compared with relatively few mutton sheep farmers (17%). 143

Over 94% of Dorper sheep farmers utilized jackal-proof fences, game fences, or electric fences in some part of their farm, compared to 23% of Karakul farmers. Less than half of Karakul farmers reported they did not have fences and the remainder reported stock fences rather than jackal-proof fences.

Unlike Dorper sheep, Karakul sheep naturally flock into one group with the young lambs in the center. They often guard their young from predators and actively defend them using their horns. Because Karakul flock, they can be herded with a shepherd and protected by guarding dogs from predators (Coppinger et al. 1998). The flock goes out each morning with a shepherd

(often accompanied with guarding dog(s)) and spend the day grazing in an area, then returning as a flock to a protected corral or ‘boma’ where they spend the night guarded by the dog. Half of

Karakul farmers reported using livestock guarding dogs while no mutton farmers utilized this strategy.

The commercial shift from a flocking breed to a non-flocking breed impacted livelihoods of local workers, because raising Dorper sheep is not conducive to using shepherds or daily management of flocks. The result was a sharp decline of employment of local people by commercial farmers. To further complicate matters, the decline in shepherd employment has resulted in a higher degree of flock theft by people, some of which were formerly employed as shepherds. Unfortunately, predators are often blamed for sheep missing due to human theft, further fueling retaliatory killing of carnivores (A. Neils, unpublished data). To offset higher rates of theft, many farmers overstock their land which amplifies another negative impact of

Dorper sheep.

144

Forage and resource use- The amount of land needed to raise meat sheep is far greater than is needed to produce pelts from Karakul. Southern Namibian commercial livestock farms must be very large to sustain themselves and make a profit. Most farms are at least 10,000 ha and some are over 20,000 ha because forage in the landscape is sparse and low in quality.

Commercial meat sheep breeds were selected to grow fast and have a large carcass weight

(Lategan 2004). They require a tremendous amount of land since they consume large quantities of plant biomass. Female mutton breeds especially need to consume a high amount of forage and water resources in order to effectively produce enough milk to nourish fast growing young.

Unlike Karakul, a Dorper lamb must be raised to 3-6 months of age (Lategan 2004). This can be incredibly taxing to the environment, particularly in drought years. Mutton sheep require a consistent output of lambs annually in order to consistently meet commercial demands.

Foraging patterns of herding sheep better suit the native landscape compared to overstocked non-flocking mutton sheep. Native grass co-evolved with herding ungulates, such as springbok, and is naturally ‘rested’ between periods of intense grazing, providing time for regeneration (Strohbach 2001). Bunchgrass grows better when intensely grazed by herbivores and rested for the rest of the season (Strohbach 2001). This removes older stems from the outside of the bunch and allows new stems to photosynthesize in the center. By replicating natural feeding on grasses, herding livestock such as Karakul can actually be beneficial to the environment.

Overstocking of mutton breeds often requires groundwater pumping to sustain bigger flocks in the arid Namibian landscape; this action is undertaken even in years of prolonged drought (A. Neils, personal observation). Pumping is not required with Karakul sheep because 145

lambs can be harvested immediately even in drought years, and flock numbers could effectively cycle up and down depending on annual rainfall.

Exposure to predators- Karakul lambs are harvested when only days old (Bravenboer

2007). Thus, young are never out on the landscape vulnerable to predation. In drought years, which are increasingly common (Quan et al. 2004), all lambs can be harvested to decrease the amount of forage that ewes need to produce milk for lambs, which is inherently already minimal.

In highly productive years, more lambs can be retained to quickly increase the herd. Stillborn

Karakul lambs, which may be seasonally common after rains when poisonous plants emerge (A.

Neils, personal observation), still have usable and valuable skins for market.

Dorper sheep tend to not flock and instead are spaced out on the landscape (Du Toit

2008). When startled, they flee in separate directions and tend not to flock; Dorper ewes abandon lambs when confronted by potential threats and do not effectively defend young (A.

Neils, personal observation). This behavior leaves lambs susceptible to predators and is the reason why guard dogs are ineffective against predators for this non-flocking breed (Coppinger et al. 1988; Smith et al. 2000).

Overall, Dorper sheep are less adapted to survive the Namibian environment, have higher need for antibiotics, higher negative impact on the landscape in terms or soil disturbance, forage less efficiently, require larger tracts of land to be used for grazing, and are more susceptible to predation resulting in higher incidences of conflict and subsequent predator control methods.

This is problematic since there is now at least 10 times the number of dorper sheep raised in

Namibian than Karakul (Table 2). Just as many individual sheep (now mutton varieties) are harvested by commercial farmers as before, they are just of a different breed and killed at an 146

older age. Whether it is better to harvest a lamb at a few days or a few months is difficult to judge. Shifting a market back to Karakul pelt production would effectively result in farmers managing flocks instead of predators. This would allow for better mitigation strategies to be implemented in response to potential carnivore conflicts (Coppinger et al. 1988; Smith et al.

2000). Shepherds and herders could be utilized again, which would also help local people find employment, reduce theft and reduce losses to predation (Shivik 2004). Flocks of Karakul could be corralled at night and guard dogs could be utilized to prevent conflicts with carnivores.

Rotational foraging could be enacted to reduce stress on the landscape and allow grazing plots to recover (Bailey and Brown 2011). This would benefit the entire landscape and a suite of native fauna that depends on using natural grassland. Reversing the market shift from raising meat specific breeds back to Karakul would benefit predators, people, and the Namibian landscape as a whole.

4.8. Conclusions

Conservation requires positive attitudes and participation of local communities

(Zimmerman et al. 2010). However HWC often causes opposition to conservation efforts

(Holmern et al. 2007). For local communities, economic impacts of coexistence, as well as perceptions and attitudes, inﬂuence tolerance of carnivores (Woodroffe & Ginsberg 1998;

Sillero-Zubiri & Laurenson 2001; Hussain 2003; Treves et al. 2006; Zimmermann et al. 2010).

Traditional and cultural anti-predator attitudes also dictate predator removal (Marker et al. 2003).

Understanding human dimensions and economics of wildlife conservation must become a central principle of carnivore conservation biology (Sillero-Zubiri & Laurenson 2001). Conﬂict

147

resolution requires customized solutions and outreach based on best available ecological data for the species involved and the social psychology of the people affected (Ajzen and Fishbein 1980).

Effective managers must have comprehensive knowledge of local livelihoods, attitudes and perceptions in order to develop successful informed management agendas that benefit all parties.

HWC management must include outreach that influences perceptions of risk (Riley and Decker

2000). Education improves predator tolerance, as does effective conflict mitigation measures

(Homern et al. 2007). A dire need exists for predator educational and extension efforts regarding the value of predators in ecosystems including farmlands, and of methods and benefits of managing farms that support game and predators. Younger farmers provide hope, as they are cognizant of these problems and more receptive to conservation collaboration and developing strategies for coexistence. Likewise, global conservation must include educating consumers about the unintended consequences of their actions and of market demands. This is equally important as educating small stock raisers who must to respond to market demand to stay in business and maintain their livelihoods. Ultimately, shifts in land use strategies are needed that integrate conservation with agricultural production. In Namibia, a large-scale return to commercial Karakul farming and/ or switch to Damama sheep for mutton production would tremendously benefit conservation of native predators across working landscapes.

148

ACKNOWLEDGEMENTS

First and foremost, the author would like to express gratitude to Timm and Inez Miller, local farming hosts Kiripotib Guest Farm (Hans and Claudia von Hase) and Tyse and Annetta Swart.

The author also acknowledges the hundreds of livestock farmers that contributed specimens, information, information, stories, and cups of rooibos tea; this research could not have done it without your cooperation and trust. Many thanks to Tom Sheridan and for his guidance and support and to John Koprowski and multiple reviewers for constructive comments. I thank

Diane Austin for her advice and guidance in my qualitative interview templates. I thank

Cheyenne Rose McChesney for her assistance transcribing and coding. This research was conducted with support from the Namibian Ministry of Environment and Tourism (under permit number 1539) and the University of Arizon’a Human Subject Protection Plan (under protocol number 1510165011). Funding was provided by Fulbright, Neils Family, M. Sloan, and contributions from many other donors. Researchers are in no way affiliated with the Namibian

Agricultural Association, karakul production, or the SWAKARA farming; no financial support came from any livestock farming association.

149

REFERENCES

Ajzen, I. and M. Fishbein. 1980. Understanding attitudes and predicting behaviour. Prentice Hall, Inc. New Jersey, USA.

Auerbach, C. and L.B. Silverstein. 2003. Qualitative data: An introduction to coding and analysis. New York University Press, New York.

Bailey, D.W. and J.R. Brown. 2011. Rotational grazing systems and livestock grazing behavior in shrub-dominated semi-arid and arid rangelands. Rangeland Ecology and Management (64): 1-9.

Barthold, J.A., A.J. Loveridge, D.W. Macdonald, C. Packer and F. Colchero. 2016. Bayesian estimates of male and female African lion mortality for future use in population management. Journal of Applied Ecology (53): 295-304.

Berger, K.M. 2006. Carnivore-livestock conflicts: effects of subsidized predator control and economic correlates on the sheep industry. Conservation Biology (20): 751–761.

Bravenboer, B. 2007 “Karakul: Gift from the Arid Land (Namibia 1907-2007),” Namibia Digital Repository, accessed 1/5/2018, http://namibia.leadr.msu.edu/items/show/157.

Cooley, H.S., R.B. Wielgus, G.M. Koehler, H.S. Robinson and B.T. Maletzke. 2009. Does hunting regulate cougar populations? A test of the compensatory mortality hypothesis. Ecology (90): 2913-2921.

Coppinger, R., L. Coppinger, G. Langeloh, L. Gettler and J. Lorenz. 1988. A decade of use of livestock guarding dogs. Proceedings of the Thirteenth Vertebrate Pest Conference (43).

Domestic Animal Diversity Information System. Food and Agriculture Organization of the United Nations. http://www.fao.org/dad-is/en. Accessed 1 December 2017.

Du Toit, D.J. 2008. The indigenous livestock of Southern Africa. South Africa.

Emberley, J.V. 1997. The Cultural Politics of Fur. Montreal and Kingston. McGill-Queen’s University Press.

Halbritter, H. J., S. T. Smallidge, J. C. Boren and S. Eaton. 2008. A Guide to Identifying Livestock Depredation. New Mexico State University Cooperative Extension Service and Range Improvement Task Force Report 77.

150

Harter, J.H. 2004. Environmental justice for whom? Class, new social movements, and the environment: A case study of Greenpeace Canada, 1971-2000. Labour/Le Travail:83- 119.

Holmern, T., J. Nyahongo and E. Røskaft. 2007. Livestock Loss Caused by Predators Outside Serengeti National Park, Tanzania. Biological Conservation (135): 518–526.

Hoogesteijn, R. 2003. Manual on the Problem of Depredation Caused by Jaguars and Pumas on Cattle Ranches. Wildlife Conservation Society, New York.

Hussain, S. 2003. The status of the snow leopard in Pakistan and its conflict with local farmers. Oryx (37): 26-33.

Kamler, J.F., P.S. Gipson and T.R. Snyder. 2000. Dispersal characteristics of young bobcats from northeastern Kansas. The Southwestern Naturalist (45): 543-546.

Knowlton, F.F. 1972. Preliminary interpretations of coyote population mechanics with some management implications. Journal of Wildlife Management (36): 369-382.

Lambert, C.M., R.B. Wielgus, H.S. Robinson, D.D. Katnik, H.S. Cruickshank, R. Clarke and J. Almack. 2006. Cougar population dynamics and viability in the Pacific Northwest. The Journal of Wildlife Management (70): 246-254.

Lategan, D. 2004. Dorpers into the New Century: Brochure and Training Manual. Dorper Sheep Breeders' Society of S.A. Middelburg, South Africa: 1-88.

Lee, M., 2014. The effects of product information on consumer attitudes and purchase intentions of fashion products made of fur, leather, and wool. Iowa State University Graduate Theses and Dissertations. Paper 13840. Linnell, J.D., J. Odden, M.E. Smith, R. Aanes and J.E. Swenson. 1999. Large carnivores that kill livestock: do “problem individuals” really exist. Wildlife Society Bulletin (27): 698-705. Loveridge, A.J., S.W. Wang, L.G. Frank and J. Seidensticker. 2010. People and wild felids: conservation of cats and management of conflicts. Biology and conservation of wild felids (161). Marker, L.L., M.G.L. Mills and D.W. Macdonald. 2003. Factors influencing perceptions of conflict and tolerance toward cheetahs on Namibian farmlands. Conservation Biology, (17): 1290-1298. Miescher, G. 2012. Namibia's red line: the history of a veterinary and settlement border. Palgrave MacMillan, New York.

151

Miquelle, D.G., E.N. Smirnov, T.W. Merrill, A.E. Myslenkov, H.B. Quigley, M.G. Hornocker and B. Schleyer. 1999. Hierarchical spatial analysis of Amur tiger relationships to habitat and prey. In: Riding the Tiger: tiger conservation in human-dominated landscapes. Cambridge University Press, Cambridge, UK, 71-99. Moolman, L.C., 1986. Aspects of the ecology and behavior of the caracal Felis caracal Schreber 1776 in the Mountain Zebra National Park and on the surrounding farms. Master’s thesis, University of Pretoria, Pretoria. Muir-Leresche, K., and R. Nelson. 2000. Private property rights to wildlife: the southern African experiment. ICER. Nowell, K., Jackson, P. 1996. Wild cats: status survey and conservation action plan. Gland: IUCN: 382. Ogada, D.L. 2014. The power of poison: pesticide poisoning of Africa's wildlife. Annals of the New York Academy of Sciences (1322): 1-20. Ogada, M.O., R. Woodroffe, N.O. Oguge and L.G. Frank. 2003. Limiting depredation by African carnivores: the role of livestock husbandry. Conservation Biology (17): 1521- 1530. Oklahoma State University. 2015. Department of Animal Science. http://www.ansi.okstate.edu/breeds/sheep Accessed 11 October 2017.

Quan, J., D. Barton and C. Conroy. 1994. A preliminary assessment of the economic impact of desertiﬁcation in Namibia. Directorate of Environmental Affairs, Windhoek. Research Discussion Paper (3): 1–148.

Rabinowitz, A.R. 1986. Ecology and behaviour of the Jaguar (Panthers onca) in Belize, Central America. Journal of Zoology (210): 149-159.

Riley, S.J.A. and D.J. Decker. 2000. Risk Perception as a Factor in Wildlife Stakeholder Acceptance Capacity for Cougars in Montana. Human Dimensions of Wildlife (5): 50–62.

Robinson, H.S., R.B. Wielgus, H.S. Cooley and S.W. Cooley. 2008. Sink populations in carnivore management: cougar demography and immigration in a hunted population. Ecological Applications (18): 1028-1037.

Rolley, R.E., 1985. Dynamics of a harvested bobcat population in Oklahoma. The Journal of Wildlife Management (49): 283-292.

Ross, P.I. and M.G. Jalkotzy. 1992. Characteristics of a hunted population of cougars in southwestern Alberta. The Journal of Wildlife Management (56): 417-426.

152

Sacks, B.N. 2005. Reproduction and Body Condition of California Coyotes (Canis latrans). Journal of Mammalogy (86): 1036-1041.

Schneider, H.P. 1994. Animal health and veterinary medicine in Namibia. Agrivet, Windhoek, Namibia.

Schoeman, S.J. 1998. Genetic and environmental factors influencing the quality of pelt traits in Karakul sheep. South African Journal of Animal Science (28): 125-139.

Sillero-Zubiri, C. and M.K. Laurenson. 2001. Interactions between carnivores and local communities: Conflict or co-existence? Conservation Biology Series- Cambridge: 282- 312.

Slough, B.G. and G. Mowat. 1996. Lynx population dynamics in an untrapped refugium. The Journal of Wildlife Management (60): 946-961.

Smith, M.E., J.D. Linnell, J. Odden and J.E. Swenson. 2000. Review of methods to reduce livestock depradation: Guardian animals. Acta Agriculturae Scandinavica, Animal Science (50): 279-290.

Stein, A. 2006. Interactions: Leopards vs. Brown Hyenas. Waterberg Carnivore Project Namibia. Otjiwarongo, Namibia.

Shivik, J.A. 2004. Non-lethal Alternatives for Predation Management. Sheep & Goat Research Journal (19): 64-71.

Strohbach, B.J. 2001. Vegetation survey of Namibia Namibia wissensschaftliche gesellschaft. Namibia Scientific Society Journal (49): 93-124.

Stuart, C.T., 1981. Notes on the mammalian carnivores of the Cape Province, South Africa. Bontebok (1): 1-58.

Stuart, C.T. and G.C. Hickman. 1991. Prey of caracal, Felis caracal, in two areas of Cape Province, South Africa. Bontebok (1): 1-58.

Stuart, C. and T. Stuart. 2013. Caracal Caracal caracal: Mammals of Africa (5): 174-179.

Sweanor, L., K.A. Logan and M.G. Hornocker. 2000. Cougar dispersal patterns, metapopulation dynamics, and conservation. Conservation Biology (14): 798–808.

Treves, A., R.B. Wallace, L. Naughton-Treves and A. Morales. 2006. Co-managing human– wildlife conflicts: a review. Human Dimensions of Wildlife (11): 383-396.

153

Woodroffe, R. and J.R. Ginsberg. 1998. Edge effects and the extinction of populations inside protected areas. Science (280): 2126-2128.

Woodroffe, R. and J.R. Ginsberg. 2005. King of beasts? Evidence for guild redundancy among large mammalian carnivores. In: Large Carnivores and the Conservation of Biodiversity. Edited by Ray, J.C., Redford, K.H., Steneck, R.S., Berger, J. Island Press: 154-175.

Wutich, A. and C. Gravlee. 2010. Water decision-makers in a desert city: Text analysis and environmental social science. In Environmental Social Sciences: Methods and Research Design. Edited by Vaccaro, E. Alden Smith, and S. Aswani. Cambridge: Cambridge University Press: 157–187

Wydeven, A.P., A. Treves, B. Brost and J.E. Wiedenhoeft. 2004. Characteristics of wolf packs in Wisconsin: identification of traits influencing depredation. People and predators: from conflict to coexistence. Island Press, Washington, DC : 28-50.

Zimmermann, A., N. Baker, C. Inskip, J.D. Linnell, S. Marchini, J. Odden, G. Rasmussen and A. Treves. 2010. Contemporary views of human–carnivore conflicts on wild rangelands. In Wild rangelands: Conserving wildlife while maintaining livestock in semi-arid ecosystems: 129-151.

154

TABLES

Table 1. Characteristics of small stock found on Namibian farmlands.

Birthin Sheep Country of Primary Drought Herding Arid Birthin Horn Management g Breed origin product tolerance instinct adaptation g s location Black Headed Somalia meat moderate low pasture moderate seasonal field no Persian Meat, Damara Egypt ** high high shepherd high seasonal corral yes hides Dorper South Africa meat moderate low pasture moderate year round field no pelt, West Karakul wool, high high shepherd high seasonal corral yes Turkestan* meat pasture/ Van Rooy South Africa meat moderate moderate moderate seasonal field no shepherd pasture/ Boer Goat South Africa meat high moderate high seasonal corral yes shepherd ** Breed isolated in Namibia *Breed selectively modified in Namibia

155

Table 2. Minimum population estimates of livestock by breed by year in Namibia.

Year 1980 1985 2014 Breed Karakul sheep 11968000 1400000 60000 Damara sheep - - 600000 Meatmaster sheep - - 2000 Black Headed Persian - - 2000 Boer Goat - - 100000 Food and Agricultural Organization of the United Nations (2017)

156

FIGURES

Figure 1. Model depicting the chain of events that followed the international anti-fur movement that led to the decline of pelt sheep in Namibia and an increase of predators removed.

Data were collected via semi-structured interviews.

157

APPENDIX D: Permits

Permits were used in this project for caracal capture and radio-collar attachment and were obtained from the following agencies:

• Namibian Ministry of Environemt and Tourism • University of Arizona’s Institutional Animal Care and Use Commeitee • Univeristy of Arionna’s Human Subject

158

Figure D-1. Initial research and collection permit issued by the Namibian Ministry of Environment and Tourism for the caracal research project.

159

Figure D-2. Initial research and collection permit issued by the University of Arizona Institutional Animal Care and Use Committee for the caracal research project.

160

Figure D-3. Social science research protocal by the University of Arizona’s Human Subjects Protection Program.

161

APPENDIX E: Caracal Necropsy Form

162

163

APPENDIX F: Farmer Qualitative Interview Template

164

165

APPENDIX G: Capture Checklist

166

APPENDIX H: Curriculum Vitae

Aletris Marie Neils

Permanent address: 3755 West Driscol Lane Office: +1 (520) 743-7686 Tucson, Arizona 85745 USA Email: [email protected]

PERSONAL

Born in Phoenix, Arizona Health: Excellent, non-smoker, 20/20 vision Flawless Criminal Record

EDUCATION

University of Arizona, School of Natural Resources, Tucson, Arizona -- (PhD - 2018) -- Wildlife Conservation and Management -- Minors in Ecology & Evolutionary Biology, Applied Anthropology, & Natural Resource Management -- Dissertation: Caracals in a Heterogeneous Landscape: Resolutions for Human- Carnivore Conflicts -- Advisors: Dr. John Koprowski & Dr. Cecil Schwalbe

University of Florida, School of Natural Resources and Environment, Gainesville, Florida -- (MSc - 2011) -- Interdisciplinary Ecology/ minor in Wildlife Conservation and Management -- Thesis: Ecology of Florida Black Bears across the Urban-Wildland Interface of Ocala National Forest -- Advisor: Dr. Melvin Sunquist

Arizona State University, Tempe, Arizona -- (BSc - 2003) -- Majored in Conservation Biology; minor in Women’s Studies -- Thesis: Western Spotted Skunk Abundance across Elevational Gradients in the Sierra Anches Mts. -- Graduated Magna Cum Laude/ Dean's list all semesters

AREAS OF INTEREST

Conservation biology; Ecology; Human-wildlife conflict resolution; Mammalogy; Environmental education

AWARDS & ACHIEVEMENTS

-- Outstanding Mentor Award (2014) -- International Collaboration Award presented from UA’s School of Natural Resources and Environment (2013) -- University of Arizona’s College of Agriculture and Natural Resources Student Award (2013) -- Rachel Carson Environmental Fellowship (2013) -- Wild Felid Association Award (2012) -- UA’s Beal Favorite Teacher of the Year Award (2012) -- UA’s Natural Resources Scholarship (2012) -- The Wildlife Society- Leadership Institute (2012) 167

-- CALS E. Zube Ecology Scholarship (2011) -- AZ Association of Environmental Professionals Scholarship (2011) -- Pat & Miriam Reid Achievement Scholarship (2011) -- Fulbright Fellowship (2010-2011) -- Science Teaching Award (2010) -- National Science Foundation BioME Teaching Fellowship (2009-2010) -- University of Florida’s Graduate Teaching Assistant Award (2007) – highest teaching honor at UF -- Agricultural Women & Vam C. York Graduate Scholarship (2006) -- Edward Abbey Young Conservationist Award (2003) -- Arizona State University’s Freshman Achievement Award (2000)

ORGANIZATIONS & PROFESSIONAL SERVICE

Society for Conservation Biology (2000- present) North American Section -- Board Member (2010- 2013) Chapter Advisory Committee -- Co- Chair & New Chapters Coordinator (2007- 2013) Madrean Chapter -- Co-Founder and President (August 2008 - January 2010) Florida Chapter -- Founder and President (April 2004 - January 2008) Central Arizona Chapter -- President (December 2000 - September 2002)

The Wildlife Society (2004- present) Women of Wildlife (WOW) member (2012- present) International Wildlife Management Working Group Board Member (2013- 2015) Leadership Institute (2012) Arizona Chapter Member (2010- present) Quiz Bowl Coach – University of Arizona (2010-2014) University of Florida Student Chapter Graduate Representative (2006) Florida Chapter Member (2004- 2008)

Large Carnivore Management Association of Namibia Member (2010- present)

Wild Felid Research and Management Association Member (2010- present)

Caracal Working Group Chair (2012- present)

Graduate Student Associations University of Arizona’s Natural Resources Graduate Association student curriculum representative (2009) University of Florida’s School of Natural Resources and Environment Graduate Association -- President (2007) -- Conservation Coordinator (2006)

168

SPECIALIZED CERTIFICATIONS

Wildland Firefighting certified (US Forest Service/ The Nature Conservancy) – 2007 Felid Anesthesia certified (University of Florida’s Veterinary School) –2007 First Aid and CPR certified – since 2002, recertified in August 2011 & 2015 Airboat and Motorboat Operator certified (US Department of Interior) – November 2007 Chemical Immobilization of Animals (Safe Capture International) – Internationally certified since 2005 Fingerprint Clearance Card – since 2009

PROFESSIONAL EXPERIENCE

Conservation CATalyst, Tucson, Arizona -- (May 2008- present) -- Position: Executive Director, Founder -- Description: Operate an NGO; fundraise; direct international feline research & education programs

Humboldt State University, Arcata, California -- (August 2017- present) -- Position: Wildlife Lecturer -- Description: Develop and instruct novel wildlife courses, labs, field trips, and seminars

Hiaki High School, Tucson, Arizona -- (October 2015 - April 2017) -- Position: STEM Advisor for Native Youth -- Description: Direct native high school students in non-traditional STEM curriculum; mentor

National Geographic Channel, Los Angeles, California -- (October 2014 - April 2015) -- Position: Expert Scientist for Everything You Didn’t Know About Animals -- Description: Translate wildlife science to the public on camera; consult on wildlife facts

Andalas University, Padang, Sumatra, Indonesia -- (September 2015 – January 2016) -- Position: Guest Instructor -- Description: Advise student research projects; mentor

University of Arizona, Tucson, Arizona -- (June 2013 – August 2015) -- Position: Mammalogist/Lecturer -- Description: Design/direct mammalogy course & trips; supervise TAs; coordinate museum specimens

University of Arizona, Tucson, Arizona -- (July 2011- 2013) -- Position: Lab/Field Instructor -- Description: Teach wildlife ecology and management lab techniques; direct field trips and labs

Salazar Hounds/ AZGFD, Mesa, Arizona -- (October 2009 - September 2011) -- Position: Contract Puma Field Biologist -- Description: Track pumas on mules with hounds-men; dart and radio collar treed cats for research

Cheetah Conservation Fund, Otjiwarongo, Namibia -- (May 2010- March 2011) -- Position: Senior Research and Education Specialist/ Conservation Biology Instructor -- Description: Instruct Conservation Biology courses for international professional participants; research

University of Florida International Center, Gainesville, Florida -- (October 2007- May 2009) -- Position: Namibia Program Director & Instructor -- Description: Develop & direct a study abroad program; teach five wildlife courses; oversee research

169

US Geological Service, Gainesville, Florida -- (May 2007- May 2009) -- Position: Biological Scientist -- Description: Write/ review reports & manuscripts; aquatic field research; isotope lab work

University of Florida, Gainesville, Florida -- (January 2005- August 2008) -- Position: Graduate Student Lab Instructor & Teaching Assistant -- Description: Teach wildlife identification and natural history, lecture, and direct field trips

University of Florida Athletic Association, Gainesville, Florida -- (August 2006- November 2007) -- Position: Science Tutor -- Description: Tutor NCAA men's basketball team and other student athletes in science and math

Florida Fish and Wildlife Conservation Commission, Gainesville, Florida -- (February 2005- April 2008) -- Position: Black Bear Biologist -- Description: Trap, immobilize and monitor Florida black bears; manage interns and volunteers

Wildlife Conservation Society, Mesoamerica Office, Gainesville, Florida -- (May 2005- May 2006) -- Position: Crocodilian Specialist -- Description: Crocodilian captures/ field research; write manuscripts/ manuals/ reports; manage accounts

Florida Fish and Wildlife Conservation Commission, Gainesville, Florida -- (September 2004- January 2006) -- Position: Small Game Check Station - Parker Field Manager -- Description: Collect data on harvested game and inform hunters on regulations; supervise volunteers

University of Florida FLREC, Davie, Florida -- (January 2004- January 2005) -- Position: Wildlife Research Biologist -- Description: Catch/ survey American crocodiles and alligators, operate boats, & write proposals/ grants

Phoenix Zoo, Phoenix, Arizona -- (March 2001- January 2004) -- Position: Program Specialist; Conservation Coordinator Assistant -- Description: Handle and present animals; write reports and recommendations for T&E AZ wildlife

Borderland Jaguars, Tucson, Arizona -- (May 2002- January 2004) -- Position: Carnivore Research Technician -- Description: Track/ monitor jaguar, puma, & bear with hair snares & camera traps; analyze hair samples

Awakening Seed School, Phoenix, Arizona -- (October 2003- January 2004) -- Position: Substitute Teacher -- Description: Teach classes from pre-school to fifth grade

Animal Planet/ Discovery Channel, King of the Jungle -- (August 2003) -- Position: Animal Expert Competitor -- Description: Perform extreme physical challenges, handle and present exotic wildlife on camera

Arizona Game and Fish, Tucson, Arizona -- (Summers 2002 & 2003) -- Position: Non-Game Field Biologist Intern -- Description: Survey vertebrates, beaver impact assessment, band raptors, and mist-net bats

US Fish and Wildlife Service, Guadalupe Mt. National Park, Texas -- (February 2003- April 2003) -- Position: Independent Contractor -- Description: Mountain lion dietary analysis in relation to potential bighorn sheep reintroduction

170

Cheetah Conservation Fund/ Round River Conservation, Namibia, Africa -- (September 2002- January 2003) -- Position: Student Research Assistant -- Description: Develop and test non-invasive methods to census felid populations; assist with monitoring Black rhinos; assist with trapping, immobilizing of felids and other wildlife

Arizona State University, Department of Biology, Tempe, Arizona -- (Summer 2003) -- Position: Mammalogy and Techniques in Conservation Biology Teacher's Assistant -- Description: Teach mammal identification, demonstrate field techniques, and oversee research projects

VOLUNTEER EXPERIENCE

Agricultural Association of Namibia, Keetmonshoop, Namibia- (2010- 2016) -- Give presentations on mitigating conflicts with predators on farmlands; depredation identification -- Recommend humane capture and handling techniques

Buenos Aries National Wildlife Refuge & Kings Anvil Ranch, AZ (2010-2016) -- Coordinate and direct annual jackrabbit surveys conducted by citizen scientists

Biosphere, Tucson, AZ – (2010-2016) -- Give presentations to the public about carnivores and other mammals during special events -- Design interactive programs on carnivore ecology

Saguaro National Park, Tucson, AZ -- (Fall 2011) -- Mammalogist/ Scientist for BioBlitz -- Capture and handle mammals and translate wildlife capture techniques to the general public

Gray Hawk Nature Center, Sierra Vista, AZ – (Spring 2010) -- Design & direct a 3 day camping trip/ ecology field course for 72- 4th to 6th graders -- Teach Sherman trapping, snare sets, mark-recapture, camera traps, mist-netting, turtle traps

Society for Conservation Biology, Gainesville, FL – (September 2007- January 2009) -- Design and coordinate a wildlife track identification, sign recognition, and data collection workshop -- Develop a long-term wildlife monitoring program to examine the effects of road expansion on wildlife

Sky Island Alliance, Tucson, Arizona -- (May 2001- January 2004; 2008-2011) -- Teach workshops on carnivore tracking & natural history; monitoring transects; GPS mapping

Northern Jaguar project, Sierra de los Pavos, Sonora, Mexico -- (Summer 2003) -- Radio telemetry triangulation of collared jaguars and pumas, track small cats and other carnivores -- Monitor camera traps

Phoenix Zoo, Phoenix, Arizona -- (January 2001- May 2003) -- Endangered black-footed ferret breeding program and husbandry -- Endangered Ramsey canyon leopard frog breeding program and reintroductions

Active Conservation & Humanitarian Activities: -- Girl Scouts of America; wildlife tracking guide; Arizona – (2015 to present) -- Low-income and homeless pet vaccination drive; Tucson -- (April 2014) -- Earth camp volunteer instructor; Sonoran Desert Museum, Tucson -- (Summer 2009) -- Directed a workshop on living with wildlife to the Boy Scouts of America; Florida -- (January 2007) -- Guided field trips & presented information on living with bears; Umatilla bear festival -- (2005, 2006) 171

-- Removed exotic invasive flora and fauna; Florida Keys, Florida -- (May 2006) -- Rehabilitation of injured wildlife & outreach on living with wildlife; Fort Lauderdale FL -- (2004) -- Tracked mountain lion and other large mammals; Fort Huachuca, Arizona -- (June 2001-2003) -- Cetacean transect surveys; Baja California, Mexico -- (March 2003) -- Seining and identification of native and exotic fish; Araivipa Canyon, Arizona -- (April 2002) -- Razor-backed sucker adult and larvae capture and tag; Lake Mojave, Nevada -- (March 2002) -- Canada goose family count; Cibola National Wildlife Refuge, California -- (January 2002) -- AZ Game & Fish Check Stations (quail, rabbit, deer, sandhill crane, javelina) – (Sept. 2001- Feb. 2002) -- Burrowing owl habitat restoration and reintroductions AZPF; Laveen, Arizona -- (November 2001) -- Black-tailed prairie dog habitat restoration and relocation; Nutt, New Mexico -- (August 2001)

ACADEMIC TEACHING EXPERIENCE

Course Role Years Institution Apex Predator Conservation & Mgmt Instructor- design class/labs/trips 2017/8 Humboldt State University Human Wildlife Conflict Instructor- design class/labs/trips 2018 Humboldt State University Wetlands Habitat Ecology Instructor- design class/labs/trips 2017 Humboldt State University Senior Seminar Instructor- design/ moderate class 2017 Humboldt State University Wildlife Biology Guest Instructor 2015 Andalas University AZ Carnivore Ecology Instructor- design class/trips 2015 University of Arizona Mammalogy Instructor- design class/labs/trips 2013/4 University of Arizona Wildlife Management Lab instructor - direct labs/ trips 2012 University of Arizona Wildlife Management for Mammals Lab instructor - direct labs/ trips 2011 University of Arizona Conservation Biology Instructor- direct classes and trips 2010/1 Cheetah Conserv. Fund Namibian Flora and Fauna Instructor- design & direct class 2008/9 University of Florida Namibian Heritage Instructor- design & direct class 2008/9 University of Florida Ecological Techniques Instructor- design & direct class 2008/9 University of Florida Wildlife Research Techniques Instructor- design & direct class 2008/9 University of Florida Wildlife Conservation & Management Instructor- design & direct class 2008/9 University of Florida Wildlife of Florida Lab instructor - direct labs/ trips 2007/8 University of Florida Wildlife Field Techniques Section instructor - direct trips 2007/8 University of Florida Wildlife Issues TA – guest lecture & support 2007/8 University of Florida Human Dimensions TA – direct labs & lecture support 2007 University of Florida Conservation Biology TA – design & direct labs 2006 University of Florida Habitat Management TA – direct field trips/ grading 2006 University of Florida Conservation Biology Field Techniques TA - direct field trips 2003 Arizona State University Mammalogy TA - design labs/direct field trips 2003 Arizona State University

PUBLICATIONS

Brown, D. E., E. Makings, A. Neils, D. Jenness, R. L. Glinski, R. D. Babb and M. B. Traphagen. 2017. Biotic Resources of the Lower Santa Cruz River Flats, Pinal County, Arizona. Desert Plants. 32(2):1-52. Neils, A. 2015. Promoting Women in Leadership Positions for Conservation of Indonesian Biodiversity. Journal of Indonesian Natural History. 3(2): 3-6. Neils, A. M., C. Bugbee, and M. Culver. 2015. Arizona Jaguar and Ocelot Conservation Education Project: K-12 Education Plan, Educational Materials, & Examples of Curricula. Final Report to the U.S. Fish and Wildlife Service under Intra-Agency Agreement Number: G13AC00222. 62pp. Neils, A. 2013. The TWS Leadership Institute. The Arizona Wildlifer. 1: 8-10. Neils, A. M. Ecology of the Florida Black Bear across the Urban- Wildland Interface of Ocala National Forest. Thesis. University of Florida, 2011. Gainesville: UMI, 2011. Print. 172

Hostetler, J., J. W. McCown, E. Garrison, A. Neils, M. Sunquist, S. Simek, and M. Oli. 2009. Demographic consequences of anthropogenic influences: Florida black bears in north-central Florida. Biological Conservation. 142(11): 2456-2463. Demopoulous, A., A. Neils, and D. Gualtieri. 2009. Synthesis of available information for Florida East and West Coasts relevant to evaluating potential environmental impacts associated with offshore sand dredging for beach and coastal restoration. USGS. Final Report. Neils, A. and C. Bugbee. 2007. Rana catesbeiana (American Bullfrog) diet. Herpetological. Review. 38(4): 443. Neils, A. 2004. Accessorized Alligators: Aquatic Reptiles Telemetry Research. Turtle River Times. 8:3.

MANUSCRIPTS IN REVIEW OR PREPARATION

Neils, A. and J. W. McCown. Florida Black Bears (Ursus americanus floridanus) at the Urban-Wildland Interface: Are They Different? The Journal of Wildlife Management. Submitted. Neils, A. and C. Bugbee. Feather- tailed Treeshrew (Ptilocercus lowii) Occurrence and Feeding Behavior in Southern Borneo. Mammalia. Submitted. Neils, A. and C. Bugbee. Bear Down! Black Bear on the Menu for American Jaguars. Frontiers in Ecology and the Environment. Submitted. Neils, A. Oral Immobilization for wild felids. Journal of Mammalogy. In preparation. M., Hemami, and A. Neils. Habitat Suitability Modeling for Caracals (Caracal caracal shmitzi) and Jebeer Gazelle (Gazella bennettii) in Abbas Abad Prohibited Hunting Area, Iran. In preparation. Irajy, F., M. Hemami, and A. Neils. A GIS-based Approach for Zoning Protected Areas: Selecting Recreation Zones in Abbas-Abad Wildlife Refuge, Central Iran. In preparation. Neils, A. How the Animal Rights Movement Indirectly Increased Predator Persecution in SW Africa. Conservation Biology. In preparation. Neils, A., and C. Bugbee. Defining Residency for Large Carnivores. Journal of Wildlife Management. In preparation. Neils, A. Feeding Ecology of Namibian Caracals. Journal of Mammalogy. In preparation. Neils, A. Namibian Aardwolf (Proteles cristata) Diet. African Journal of Ecology. In preparation.

SELECTED PRESENTATIONS

Neils, A. 2016. Becoming a Naturalist. Presented to Yale School of Forestry and Environment. Yale. New Haven, CT. November 2016. Neils, A., and C. Bugbee. 2016. Jaguar Conservation in America. Presented to the New England Conservation Association. Old Lyme, CT. November 2016. Neils, A., and C. Bugbee. 2016. Jaguar Conservation in America. Presented to the Biosphere 2’s Distinguished Researchers Symposium. Oracle, Arizona. April 2016. Neils, A. 2015. Conservation Strategies for Mitigating Conflicts between Predators and People. Presented at International Wildlife Symposium. Padang, Sumatra, Indonesia. 3 November 2015. (Keynote Speaker) C., Bugbee, and A. Neils. 2015. American Jaguar Conservation. Presented at International Wildlife Symposium. Padang, Sumatra, Indonesia. 3 November 2015. Neils, A., R. Babb, D. Brown, and C. Bugbee. 2013. Status and Distribution of the Opossum (Didelphis virginiana) in Arizona. Presented to the Arizona/ New Mexico Wildlife Society Conference. Pinetop, Arizona. February 2014. Neils, A., M. Cook, and C. Ayers. 2012. Increasing Diversity among Underrepresented Demographics in the Wildlife Profession. Presented at the Wildlife Society Meeting. Portland, Oregon. July 2012. Neils, A. 2012. The Madrean Chapter of SCB. Presented at the NA SCB Congress. Oakland, California. July 2012. Neils, A. 2012. Caracal Conservation in a Heterogeneous Landscape. Presented at the International Wildlife Management Congress. Durban, South Africa. June 2012. 173

Neils, A. 2011. Caracal Conservation. Presented at the Wildlife Society Conference. Hawaii, USA. October 2011. Neils, A. and F. Nagle. 2010. Capacity Building for Society for Conservation Biology Chapters and Conservation Organizations. Presented at the Global Congress for Conservation Biology. Edmonton, Alberta, Canada. July 2010. Neils, A. 2010. Data Gaps in Caracal (Caracal caracal) and Serval (Leptailurus serval) Natural History. Presented at the Namibian Wildlife Meeting, Otjiwarongo, Namibia. June 2010. Neils, A. 2010. Feasibility of Re-establishing Grizzly Bears (Ursus arctos) into the Southwest. Presented at the Wildlife Society SW Meeting, Flagstaff, AZ. February 2010. Neils, A. 2009. The Ecological Role of Spots in Felids. Presented at the Conway Ecology Symposium, Tucson, AZ. November 2009. Neils, A. 2009. The Role of Chapters in the Society for Conservation Biology. Presented at the Global Congress for Conservation Biology. Beijing, China. July 2009. Neils, A. 2008. Global Chapters of the Society for Conservation Biology. Presented at the Colorado Plateau Chapter of the Society for Conservation Biology Meeting. Lees Ferry, AZ. October 2008. Neils, A. 2008. Getting the Adult Community Involved in Conserving Local Wildlife: A Case Study from Florida. Presented at the Society for Conservation Biology Meeting. Chattanooga, TN. July 2008 Neils, A. et al., 2007. Ecology of the Florida Black Bear across the Urban Wildlife Interface of ONF. Presented at the Society for Conservation Biology Meeting. Port Elizabeth, South Africa. July 2007. Neils, A. et al., 2006. Ecology of the Florida Black Bear across the Urban Wildlife Interface of Ocala National Forest. Presented at the Carnivores Conference. St. Petersburg, Florida. November 2006. Neils, A. et al., 2005. Florida Black Bear Ecology and Conservation. Presented at the Cooperative Committee Meeting. Gainesville, Florida. March 2005.

PROFESSIONAL REVIEWS PROVIDED

Journals Conservation Biology; Journal of Indonesian Natural History Books Hunter, L. 2015. Wild Cats of the World. Bloomsbury Natural History. 240 pages. Reports Society for Conservation Biology; US Fish & Wildlife Service; Defenders of Wildlife; Arizona Game & Fish Department

ADVISED STUDENTS

Catherine C. Cavanaugh, Co-advised for Honors Thesis Titled: Perceptions of urban bobcats by residents of the Tucson area: Assessing the effectiveness of education and community outreach. U of Arizona. May 2013.

Selected Advised Student Research Presentations Ethan Sandoval, Kristin Ulvestad, Julia Muldoon and Aletris Neils. Assessment of Biodiversity in the Northern Jaguar Reserve in Sonora, Mexico and its Influence on the Jaguar (Panthera onca). Presented to the Arizona/New Mexico Wildlife Society Conference. Las Cruces, New Mexico. February 2015. Jason Myrand, Lisa Lang, David E. Brown, Randy Babb, and Aletris Neils. Impacts of Natural and Anthropogenic Influences on Jackrabbit Populations in Desert Grasslands of Southern Arizona. Presented to the Arizona/New Mexico Wildlife Society Conference. Las Cruces, New Mexico. February 2015. Jennifer Merems and Aletris Neils. Small Mammal Communities Response to Environment via Barn Owl pellets in Namibia. Presented to the Arizona/New Mexico Wildlife Society Conference. Las Cruces, New Mexico. February 2015.

174

Riley Bogart, Melissa Stevenson, Robert Fink, and Aletris Neils. Supplemental Water Methodology for Reintroduced Black-tailed Prairie Dog Colony. Presented to the Arizona/New Mexico Wildlife Society Conference. Las Cruces, New Mexico. February 2015. Nerissa Hall, Jennifer Merems, and Aletris Neils. Namibian Small Mammal Research via Barn Owl (Tyto alba) Pellets. Presented to the Arizona/ New Mexico Wildlife Society Conference. Pinetop, Arizona. February 2014. Levi Heffelfinger, Megan Hurrell, Dave Brown, Jim Heffelfinger, and Aletris Neils. Cottontail Rabbit (Sylvilagus audubonii) Abundance Trend in Southeastern Arizona. Presented to the Arizona/ New Mexico Wildlife Society Conference. Pinetop, Arizona. February 2014. Christina Lauren Erdelyi, Michael James Taylor, Theresa Nicole Huckleberry, David E. Brown, Randall D. Babb, and Aletris Neils. Effectiveness of Mitigation Strategies on Alfalfa Grazing by Black-Tailed Jackrabbits and Desert Cottontails in Chandler’s Veterans Oasis Park. Presented to the Arizona/ New Mexico Wildlife Society Conference. Pinetop, Arizona. February 2014. Tayler LaSharr, Sarah Schwenck, Christopher Bugbee, and Aletris Neils. Reaction of Coyote (Canis latrans) to a Variety of Olfactory Stimuli from Interspecific Carnivores. Presented to the Arizona/ New Mexico Wildlife Society Conference. Pinetop, Arizona. February 2014. *Honored as the top student presentation

Selected Advised Student Research Assists & Interns Ethan Sanderval Jaguar Camera Trap Analysis University of Arizona Spring 2015 Katlin Martin Jaguar Education Curriculum University of Arizona Fall 2014 Jennifer Merems Namibian Owl Diet University of Arizona Fall 2014 Tayler LaSharr AZ Mammalian Surveys University of Arizona Fall 2014 Amanda Veals Namibian Carnivore Research University of Arizona Summer 2013 Colleen Cavanaugh Namibian Carnivore Research University of Arizona Summer 2013 Elan Prince Henry Fox Namibian Carnivore Research University of Arizona Summer 2012 Lily Harrison Namibian Carnivore Research University of Montana Summer 2012 Allison Pena Namibian Carnivore Research University of Arizona Summer 2012 Samantha Jones Namibian Carnivore Research University of Bristol Summer 2011 Sean Thoman Namibian Carnivore Research Arizona State University Summer 2011 Elin Gromberg Human-wildlife Conflict Mitigation Swedish U. Agricultural Science Summer 2011 Magnus Nystrand Human-wildlife Conflict Mitigation Swedish U. Agricultural Science Summer 2011 Jessica Ugararte Carnivore Conflict Mitigation Polytechnic of Namibia 2010-2011 Sarah Purcell Black Bear Cub Ecology University of Florida 2007-2008 Brian Smith Black Bear Ecology University of Florida 2006-2008 Dan Gualtieri Black Bear Ecology University of Florida 2006-2008 Cameron Kovach Black Bear Ecology University of Florida 2005-2008 Katherine Issacks Black Bear Telemetry University of Florida 2005-2007

SELECTED RESEARCH FEATURED IN THE MEDIA

In Print bioGraphic, September 2017, “Up Against the Wall” by Julian Smith. Cover Story: http://www.biographic.com/posts/sto/up-against-the-wall Sierra Magazine, August 2017, “How Trump's Border Wall Could Block the Most Exciting Wildlife Comeback in North America” by: Jason Mark. Cover Story. Pacific Standard, August 2017, “Cat Fight: Inside the Struggle to Save the Latest Migrants Crossing the U.S.- Mexico Border” by: Alan Wiseman. Cover Story. Smithsonian, October 2016, “The Return of the Great American Jaguar” by: Richard Grant. Cover Story. The Bark, April 2016, “On the Jaguar’s Trail” by: Karen B. London. Cover Story.

175

Cat Fancy Magazine, September 2013, “The Most Hunted Cat in the World: A Little-known Cat Species, the Caracal, Faces Numerous Threats” by: Brad Kollus. National Geographic Kids Magazine, April 2013, “The Secret Life of the Caracal.” By: Alexander Braczkowski. Agriforum Magazine, April 2013, Namibian Farmers of the Year Award. Agriforum Magazine, May 2011, Rooikats. The Namibian, Newspaper. First Caracal Study Underway. FrontPage. 21 July 2011. New York Times, 2008, Sturgeon Research. FrontPage.

On screen/audio Discovery Channel. Daily Planet. “Arizona Jaguar Research” June 2016. National Geographic Channel. Everything You Didn’t Know About Animals. “Lions, Owls & Frogs”. Season 1: Episode 2. 20 April 2015. National Geographic Channel. Everything You Didn’t Know About Animals. “Tigers, Rhinos & Naked Mole Rats”. Season 1: Episode 5. 11 May 2015. National Geographic Channel. Everything You Didn’t Know About Animals. “Orangs, Raccoons, and Cool Cats”. Season 1: Episode 6. 18 May 2015. Mrs. Green’s World. Podcast aired 24 March 2014. http://www.mrsgreensworld.com/2014/03/24/carson-scholar- aletris-neils/ Namibian Public Radio, July 2011 “Caracal research on Farmlands” Incredible Scientists. 11 January 2011. https://www.youtube.com/watch?v=tY_K1D6xSu8 Animal Planet. King of the Jungle. Season 1: Episodes 1-9. 2003, re-aired in 2004.

Online USA Today, September 2017 “Endangered Species, and the Wall that Could Silence Them”,: https://www.usatoday.com/border-wall/story/environmental-impact-endangered-species-jaguar/582602001/ Earth Touch News Network, 19 October 2016, “America's celebrity jaguar 'El Jefe' is a bear hunter”: https://www.earthtouchnews.com/natural-world/predator-vs-prey/americas-celebrity-jaguar-el-jefe-is-a- bear-hunter/ LymeLine, 29 September 2016, “Lyme-Old Lyme HS Alum’s Work Tracking Only Wild Jaguar in US”: http://lymeline.com/2016/09/lyme-old-lyme-hs-alums-work-tracking-only-wild-jaguar-in-us-featured-in- current-smithsonian-magazine/ Univision, 12 April 2016, “Hay más tigres en cautiverio en Estados Unidos que libres en todo el mundo”: http://www.univision.com/noticias/planeta/hay-mas-tigres-en-cautiverio-en-estados-unidos-que-libres-en- todo-el-mundo Live Science, 3 March 2016, “Arizona's Only Jaguar Prowls a Difficult, But Hopeful, Path”: https://www.livescience.com/53892-arizona-only-jaguar-prowls-difficult-path.html New Times, 4 February 2016, “Only Known Wild Jaguar Spotted by Wildlife Camera in Southern Arizona”: http://www.phoenixnewtimes.com/news/only-known-wild-jaguar-spotted-by-wildlife-camera-in-southern- arizona-8025972 Discovery News, 4 February 2016, “America's Only Wild Jaguar Caught on Video”: http://www.seeker.com/americas-only-wild-jaguar-caught-on-video-1770839552.html Los Angeles Times, 3 February 2016, “He roams alone: El Jefe may be the last wild jaguar in the U.S.”: http://www.latimes.com/nation/la-na-sej-last-wild-jaguar-in-the-u-s-20160203-story.html VOA, 4 February 2016, “Only Known US Jaguar Shown in Video Roaming Arizona Mountains”: http://www.voanews.com/a/only-known-us-jaguar-shown-in-video-roaming-arizona- mountains/3176265.html ABC News, 3 February 2016, “El Jefe, only wild jaguar in US, captured on video in Arizona”: http://www.abc.net.au/news/2016-02-04/footage-captures-wild-jaguar-in-arizona-el-jefe/7139760 BuzzFeed News, 3 February 2016, “The Only Known Jaguar In The U.S. Is Roaming Around Arizona”: https://www.buzzfeed.com/jasonwells/call-me-jason-the-jaguar?utm_term=.dmPOwMXmK#.on2gvLw4b

176

Fox News, 3 February 2016, “America's only wild jaguar caught on video”: http://www.foxnews.com/science/2016/02/03/americas-only-wild-jaguar-caught-on-video.html The Christian Science Monitor, 3 February 2016, “El Jefe: Is this the United States' last jaguar?”: http://www.csmonitor.com/Environment/2016/0203/El-Jefe-Is-this-the-United-States-last-jaguar Tucson Sentinel, 3 February 2016, “Jaguar prowls Santa Ritas south of Tucson”: http://www.tucsonsentinel.com/local/report/020316_jaguar/watch-jaguar-prowls-santa-ritas-south-tucson- new-video/ Tucson News Now, 3 February 2016,”Sole jaguar in America caught on camera in Arizona“: http://www.tucsonnewsnow.com/story/31135726/sole-jaguar-in-america-caught-on-camera-in-arizona Tucson.com, 3 February 2016, “Only wild jaguar in US captured on video south of Tucson”: http://tucson.com/news/only-wild-jaguar-in-us-captured-on-video-south-of/article_599dc320-8346-5e5a- 8238-b316cf0d51e4.html Santa Cruz IMC, 3 February 2016, “New Video Shows America's Only Known Wild Jaguar”: https://www.indybay.org/newsitems/2016/02/03/18782575.php Scientific American, 5 March 2012, “The Caracal Cat Conservation Documentary Project”: http://blogs.scientificamerican.com/guest-blog/the-caracal-cat-conservation-documentary-project/

PUBLISHED WILDLIFE PHOTOGRAPHY

Publishing Body Date Smithsonian October 2016 AZ Daily Star September 2016 Discovery Channel (Daily Planet) June 2016 National Public Radio (online) February 2016 Journal of Indonesian Natural History December 2015 National Geographic Channel May 2015, re-aired September 2015 Sonorensis Magazine Winter 2013 (Volume 33; Number 1) Cat Fancy Magazine September 2013 Agriforum Magazine June 2013 National Geographic Kids April 2013 The Namibian Newspaper May 2013, July 2011

FIELD RESEARCH PROFICIENCY

Habitat types/ecoregions Sonoran/ Mohave/ Chihuahuan/ Great Basin Deserts Riparian systems of the southwestern U.S. Primary and secondary forests of the southwest and southeast Sawgrass marshes, sloughs, cypress swamps, mangrove swamps, hammocks, pinelands across the southeastern US Kalahari grasslands, sparse shrublands, and acacia woodlands across southern Africa Tropical/ subtropical moist broadleaf forests, grasslands, coniferous forests, mangroves of Sundaland bioregion Tropical/ subtropical moist and dry broadleaf forests of the Wallacea bioregion Coastal communities of the Pacific, Atlantic, Indian Oceans

Taxa: techniques Larger mammals: cage trap, snare, free-dart, immobilize, radio-collaring, camera-trap, spoor track, triangulation, aerial radio-tracking, pellet/scat counts, scent stations, distance sampling, genetic sampling, mark- resight/recapture, track plates, observation, necropsy, safely handle/ ID/ collect morphometrics Small mammals: Sherman trap, Havahart trap, Dipidomys trap, mark-recapture, fluorescent dye labeling, safely handle/ ID/ collect morphometrics 177

Bats: mist-netting, harp trap, physiological studies, radio telemetry, safely handle/ ID/ collect morphometrics Cetaceans: distance sampling, scat collection, necropsy Birds: point counts, mist netting, rocket net, ground trap, nest searches, owl call counts, safely handle/ ID/ collect morphometrics Crocodilians: noose capture, hand grab, mark-recapture, genetic sample, nest surveys, distance sampling, spotlight surveys, safely handle/ ID/ collect morphometrics Reptiles: field capture, identification, snake boards, PIT tags, turtle traps, drift fence/ pitfall traps, numeration, safely handle/ ID/ collect morphometrics Amphibians: Moller dredging, PVC pipe surveys, frog call counts, salamander surveys, crayfish trap, safely handle/ ID/ collect morphometrics Fishes: seining, electrosampling, seining, netting, tag, acoustic telemetry, snorkel/ SCUBA transect surveys, minnow trap, safely handle/ ID/ collect morphometrics Invertebrates: tree shaking/beating, enumeration of mounds, pooters, pitfall traps, sticky trap, moth cloth, field collecting Plants: point-intercept, gap-intercept, line-intercept, Daubenmire, biomass clipping Trees: measurements (DBH, height, slope, distance, crown, timber, etc.), plot and downed/ dead sampling Soils: field classification, penetrometer, hydrometer Geological: clinometer, orientation (dipping plane, linear feature, etc.), GPS, altimeter, field classification

TRAVEL

Namibia, Zambia, Zimbabwe, Botswana, South Africa, Mexico, Canada, Indonesia, Malaysia, Philippines, Bahamas, Haiti, France, United Kingdom, China, Belize

OUTSIDE INTERESTS AND ACTIVITIES

Interests: Tracking, photography, ceramics, organic gardening, SCUBA, kayaking, cooking, camping Dance: Ballet, modern, jazz, cheer, Argentine tango, salsa, ballroom, and country (rhythm & progressive) Languages: English (native); French (6 years); Afrikaans (3 years); Spanish (1 year); Cree (1/2 year)

REFERENCES

Professional References -- Professor David E. Brown Wildlife Professor & Big Game Manager University of Arizona & Arizona Game & Fish Department (retired) Phoenix, AZ USA +1 (602) 471-2872 [email protected]

-- Dr. Andrew Smith Distinguished Professor School of Life Sciences Arizona State University Tempe, Arizona 85287 USA +1 (480) 965-4024 [email protected]

178

VITA

Aletris Marie Neils was born and raised on a pecan farm outside of Phoenix, AZ where she spent her childhood with her sisters outdoors collecting and studying reptiles, insects, and other creatures that shared the land. Her conservation ethic was further nurtured during her summers on the Rocky Boy Reservation in Montana, where she learned important lessons in natural history including how to track wildlife.

She attended Arizona State University and obtained a BSc in Conservation Biology with a double minor in Women’s Studies and Performing Dance in 2003. While in school, she worked as an education specialist for the Phoenix Zoo. She soon advanced to the Conservation

Department, where she worked with many endangered species, including in the black-footed ferret breeding facility. In addition, Alertis became involved tracking pumas and jaguars in southern

Arizona and Northern Mexico with the Borderland & Northern Jaguar Project and advanced non- invasive methods for censusing large felids. In 2002, she was invited by the Cheetah Conservation

Fund to test those methods on cheetahs and leopards in Namibia.

Upon her return to the U.S., she was offered a position with the Florida Coop Unit working on crocodiles and alligators in south Florida and Belize. Aletris eventually transitioned to work with the Florida Fish and Wildlife Conservation Commission (FFWCC) Carnivore Research

Team, where she researched Florida black bears living at the urban- wildland interface. While working for FWC, she enrolled at the University of Florida and radio collared 50 bears for her

Master’s research (in Interdisciplinary Ecology). Aletris’ experiences in Florida made her realize

179

that her primary passion lies in resolving human- carnivore conflicts, especially on species without adequate ecological or behavioral data.

In 2008 Aletris founded Conservation CATalyst, a nonprofit organization dedicated to research, education, and conservation pertaining to the world’s cats (and other carnivores) that are in perpetual conflict with people. Working in Arizona (on jaguars and ocelots), southeast Asia (on leopard cats and leopards), and southern Africa (on leopards, cheetahs, and caracals), Aletris researched ecology of mammalian predators, particularly relating to species diversity and top- down ecosystem regulation. She uses this information to help formulate effective conflict resolution strategies. Improving livelihoods for local people and the fate of the predators they coexist with.

Since 2002, Aletris has worked in Namibia conducting research on caracals, servals, cheetahs, leopards, black-backed jackals, honey badgers, and African wild cats. She received a

Fulbright fellowship in 2010 to conduct her Ph.D. research (Wildlife and Fisheries Management, at the University of Arizona) focusing on ecology and conservation of caracals and other small felids on Namibian farmlands. She works closely with Namibian farmers examining perceptions of carnivores, traditional ecological knowledge, and dynamics of livelihood-specific conflicts with native predators. Aletris is collaboratively developing locally acceptable solutions to reduce carnivore conflicts and mitigate livestock depredation. She shares these strategies to other parts of the world that face similar scenarios with predators and people.

When not chasing fangs and claws, Aletris enjoys gardening, hiking, cooking, dancing,

SCUBA diving, and farming heritage chicken breeds. Aletris resides on a small ranch just outside

180

Tucson, Arizona with her husband Christopher David Bugbee, their beautiful daughter Aurelia

Lillian Lou and a menagerie of animal companions.

181