CHAPTER 6 People and wild felids: conservation of cats and management of conflicts

/Andrew J. Loveridge, Sonam W. Wang, Laurence G. Frank, and John Seidensticker

A lioness killed in an illegal wire snare by poachers. ©. AJ. Loveridge.

Introduction extirpated felid populations and still threaten many more. The ways in which people value and interact Wild felids and people have a complex and often with organisms and their habitats is at the heart of paradoxical relationship. On the one hand, human- conservation. This chapter explores some of the inter- kind admires and reveres felids; cats appear as cultural relationships between people and wild felids, where icons and symbols across the ages. In addition, there is human actions threaten felid populations. a growing awareness of the value of wild felids as key components of ecosystems, tourist attractions gener- ating income, umbrella species for conserving ecosys- tems, and flagships for engendering public support for conservation. These positive values sometimes con- Why conserve wild felids? trast strongly with the relationship between wild felids We preserve carnivores for aesthetic, symbolic, spiri- and people in areas where they coexist. Human con- tual, ethical, utilitarian, and ecological reasons. Felids flicts with wild cats, overexploitation of felid and prey are culturally valued and are important as cultural populations, and habitat loss and fragmentation have icons and symbols. Felids are widely depicted in art, 162 Biology and Conservation of Wild Felids from Stone Age petroglyphs and cave paintings to biodiversity. Felids are economic assets and, when more modern depictions of cats as art, reminding us used sustainably through tourism, trophy hunting, that humans have interacted with felids for as long as or commercial exploitation can contribute substan- we have been humans. Human culture is enriched by tially to both their own conservation and that of the symbolic use of felid images. Felids adorn many their habitats. For example, African lions (Panthera currencies, are heraldic symbols on coats of arms and led) earned Amboseli National Park, Kenya, US appear widely on the badges of national sports teams. $27,000 per lion per year in tourism revenues (West- They were revered by many cultures, for instance, the ern and Henry 1979). Trapping of furbearers or un- lion-headed goddess Sekhmet represented war and regulated trophy hunting can lead to controversial disease in ancient Egypt. The jaguar (Panthera onca) declines in populations. However, where it is carried features significantly in Central American cultures. out on a well-managed and sustainable basis, it can Similarly, the tiger (P. tigris) is culturally important provide incentives for long-term protection, a good in South and East Asian cultures (Weber and Rabino- example being sustainable harvests of Canada lynx witz 1996). (Lynx canadensis) furs. Finally, felids play an impor- Perhaps because of their place in our consciousness tant and often regulatory role within the ecosystems and the ease with which they are recognized, felids are they inhabit (Mills and Biggs 1993; Karanth and used as umbrella and flagship species to conserve hab- Stith 1999; Grange and Duncan 2006). This ecologi- itat, benefiting both felids and biodiversity generally. cal role needs to be factored into any valuation we Large carnivores are often highly mobile and viable make of felids in the wild. It must also be included in populations require large tracts of suitable habitat any assessment we make in valuing our relationship and adequate prey populations. Thus, strategies for with nature and the pristine habitats that should the protection of large felids also offer protection for form the baseline for all conservation efforts. large 'functioning ecosystems' (Soule and Simberloff 1986; Noss etal. 1996; Seidensticker era/. 1999). It is much easier to motivate the public and governments to protect a charismatic carnivore than a species that How people impact felids: anthropogenic is seldom seen and outwardly unremarkable. Exam- ples of this include the protection, through an initia- threats to felid populations tive called Paseo Panthera (the Path of the Panther), of Habitat loss is a global phenomenon, affecting all biological corridors on the Panamanian Isthmus, used species. The earth's human population has increased by jaguars, pumas (Puma concolor), and other wildlife. from 3 billion to 6 billion since 1960. The global 'Tiger Reserves' protect significant areas of biodiversi- economy has increased sixfold and food production ty in India, as well as halting the decline of tigers, and increased by 2.5 times. Nearly 25% of the earth's tigers are used as a flagship to promote conservation terrestrial surface is now under cultivation. The past efforts along border regions in the Indian subconti- 40-50 years have seen a sharp increase in the nent and Indochina (Weber and Rabinowitz 1996; amount of land converted to agriculture, and projec- Rabinowitz 1999; Seidensticker et al., Chapter 12, tions suggest that further conversion is to be this volume; Sunquist and Sunquist 2002). expected in the future (Millennium Ecosystem As- Moral philosophers and animal welfarists are sessment 2005). Conversion of natural habitat to united in their arguments that every species has an agricultural land, urban development, and destruc- intrinsic conservation value that implies it has a tion or fragmentation of habitats through logging, right to survive. Intrinsic conservation values have building infrastructure (e.g. dams, roads, power lines, served as an impetus for conservation. However, in- oil, and mineral extraction), and other human activ- trinsic value in itself has been insufficient to secure ity has serious impacts on wild felid populations. successful conservation because humans are moti- Equally important is loss of prey populations vated more by economic self-interest than by ideas through over-hunting, retaliatory persecution, habi- (Kellert et al. 1996). Economic benefits can provide tat loss, or fragmentation. Predator population den- powerful incentives to local communities to protect sities are closely correlated with prey-population People and wild felids: conservation of cats and management of conflicts 163 biomass, thus loss of prey species (either through can also be exposed to diseases carried by domestic overexploitation, such as the bushmeat trade, or carnivores, often as a result of human encroachment habitat destruction) causes linked declines in felid into wild habitats. In some populations, anthropo- populations (Karanth etal. 2004c; Henschel 2007). genic mortality can be extremely high and lead to Against a backdrop of global habitat loss, felids population declines. For instance in Laikipia, Kenya, face a number of proximate anthropogenic threats. 17 of 18 tagged lions which died were killed in retri- One of the most important ways in which people bution for livestock raiding, with the population impact felids is through increasing rates of non-nat- declining by about 4% per annum (Woodroffe and ural mortality (Fig. 6.1). For felid populations below Frank 2005). Similarly, lion density was significantly their carrying capacity, anthropogenic mortality is reduced by conflict with local people on group thought to be largely additive rather than compen- ranches surrounding Masai Mara Reserve, Kenya satory to natural levels of mortality (Lindzey et al. (Ogutu et al. 2005). In a study of Amur Tigers, all 1992). By contrast, in highly protected populations seven recorded deaths of individually recognized anthropogenic mortality is rare, for instance only adult females were due to poaching and 57% of one (7%) of 14 leopard (Panthera pardus) deaths re- cub mortality in this population was anthropogenic corded in Kruger National Park, South Africa, by (Kerleyefa/. 2002, 2003). Bailey (1993) was due to poaching; the rest were Utilization of felid populations varies substantially due to intraspecific fights, predation, and starvation. in intensity, with mortalities from this source ranging In general, anthropogenic mortality in many studied from around 7% in Eurasian lynx (Lynx lynx) in Swit- felid populations is high. While this may in part be zerland (Schmidt-Posthaus etal. 2002a) to 27 (62.7%) due to the fact that conservation biologists choose to of 43 deaths of radio-collared lions around Hwange study populations that are under threat, it is also an National Park, Zimbabwe (Loveridge etal., Chapter 11, indicator of the impact that humans have on felid this volume). Trophy hunting made up between populations. Common sources of anthropogenic 11.7% and 50% of recorded mortality in pumas mortality are legal hunting and trapping, poaching, (Logan etal. 1984; Lindzey etal. 1988; Cunningham problem animal control (both legal and illegal), and etal. 2001), 37.5% in radio-tagged leopards in Natal, vehicle accidents. Furthermore, felid populations South Africa (Balme and Hunter 2004), and trapping

Figure 6.1 Lioness killed by a steel wire snare. Wire snares set for both predators and prey species may increase levels of mortality within predator populations. (Photograph courtesy of P. Lindsey and Sango Ranch, Save Valley Conservancy.) 164 Biology and Conservation of Wild Felids made up around 45-66% of mortality of marked In each section, we offer a synthesis of the potential bobcats (L. rufus) in Mississippi and Maine, United conservation and management solutions. States (Litvaitis etal. 1987; Chamberlain etal. 1999). Trapping mortalities made up 19 (70%) of 27 recorded deaths of Canada lynx (I. amadensis) in the Northwest Territories (Poole 1994). However, for many exploited Conflicts between felids and people populations few demographic data are available and Key areas of conflict between people and felids are this is particularly true of some populations hunted through depredation on domesticated animals or and trapped for the fur trade (e.g. spotted cats in South game species and, less frequently, when large felids America in the 1960s and 1970s). kill or injure people. Livestock raiding or human Conflict with people can lead to high levels of deaths often lead to retaliatory killing of the felids mortality, particularly where predator eradication responsible. We will discuss, in the sections below, occurs. For instance, an average of 29 (range 10-42) the issues of livestock depredation and human kill- lions are destroyed each year in farmland around ing, the consequences for felid populations, and po- Etosha National Park, Nambia, where they pose a tential management solutions in situations where threat to livestock, with 563 lions killed over a such depredations occur. 20-year period (Slander 2005b). In puma populations monitored in Arizona and Utah, United States, 47% and 24%, respectively of marked animals that died were killed in defence of livestock (Lindzey et al. Consequences of conflict for felid 1988; Cunningham et al. 2001). Around 3500 pumas per year are killed by people (through sport- populations hunting, protection of livestock, and vehicle acci- Loss of human life or livelihood provides the impe- dents) in the western United States (Papouchis 2004). tus for people to attempt localized or sometimes Poaching can contribute substantially to mortality wide-scale eradication of predators. Historically, within a population. Ferreras et al. (1992) found that eradication of carnivores has been a state-supported 42% of deaths of Iberian lynx (Lynx pardinus) in Do- priority, incentivized by rewards and bounties. From nana National Park, Spain, were due to illegal poach- 1860 to 1875, 4708 tigers and leopards were reported ing. Poaching accounts for a significant proportion of killed in India (Boomgaard 2001). In the United tiger mortality in India and Russia (Kumar and Wright States, over 1160 pumas were culled from 1987 to 1999; Miquelle etal. 2005a) and Kenney etal. (1994, 1990 in an attempt to limit livestock losses in ranch- 1995) suggest that even moderate levels of poaching ing areas (Johnson et al. 2001b). Wide-scale lethal over relatively short time periods (~6 years) can lead to control of predators still occurs in some frontier massive population declines of up to 95% in this spe- areas. For example, Michalski et al. (2006a) found cies. In the Russian Far East, Chapron et al. (2008b) that 110-150 jaguars and pumas were killed over a suggest that tiger populations cannot recover if annual 12-month period by professional predator hunters in mortality rates exceed 15%. Finally, road accidents can southern Amazonia, Brazil. also be significant sources of mortality in some felid Contemporary government policies regarding pred- populations. Forty-five per cent of ocelot (Leopardus ator management are generally more enlightened. pardalis) mortality in Texas (Haines et al. 2005) and However, in areas where depredations are perceived 48% of recorded puma deaths in Florida were due to to impact livelihoods, both legal and illegal retaliatory road accidents (Taylor etal. 2002). killing of felids can extirpate populations. Frank et al. In this chapter we identify two key areas where felid (2006) describe the decimation of the lion population populations face anthropogenic threats. Firstly, we over large areas of Kenyan Masailand, where lion pop- explore the way felids impact people's lives and liveli- ulations had previously been secure. The increasingly hoods and in turn the way in which people retaliate. negative attitudes to wild predators in Masailand are We then consider the impacts of trade and overexploi- at least nominally due to livestock depredation. How- tation on the conservation status of felid populations. ever, limited involvement by local people in wildlife People and wild felids: conservation of cats and management of conflicts 165 tourism and lack of access to the revenue generated, economically significant losses to domestic livestock, coupled with ready access to agricultural poisons, may although species such as serval (Leptailurus served), wild- also be motivations behind increasing levels of illegal cat (Fells silvestris), and guina (Leopardus guigna) may predator control. In Kenya, today, the use of agricul- occasionally prey on small stock or poultry (Sanderson tural insecticide Furadan (carbofuran) to kill predators et al. 2002c; Sunquist and Sunquist 2002). Generally, is probably the single greatest source of lion mortality; species ranging in size from the caracal (Caracal caracal; in Laikipia and Kajiado districts alone, a minimum of ~12 kg) to the size of the tiger (~235 kg) have been 70 lions are known to have been poisoned since 2001 found to be the most problematic stock raiders. With (L.G. Frank, S. Maclennan, and A. Cotterill, unpub- the exception of the caracal and Eurasian lynx (~23 lished data) and populations of vultures and other kg) all of the most import livestock-killing felids are 50 scavenging birds are plummeting throughout the kg or heavier (Inskip and Zimmermann 2009). Table country (S. Thomsett, personal communication). 6.1 provides some examples of livestock depredation by felids. Comparison of trends between studies is complicated by geographic and ecological differ- ences, varying spatial and temporal scales of the stud- Predation by felids on ungulate game ies, and reporting of widely different parameters and species data. This makes rigorous evaluation of trends diffi- cult; however, there are a number of broad patterns Large felids can come into conflict with people over that emerge and these are discussed in more detail in competition for wild prey species. Predators were even the following text. persecuted as vermin in some national parks until as late as the 1950s, because carnivores were thought to suppress 'game' numbers (Davison 1967; Smuts 1976). Patterns of livestock depredation A worldwide meta-analysis of human-predator con- Different size classes of livestock are vulnerable to flict found that predators are reported to kill from different suites of felid predators. Smaller felids, such 0.02% to 2.6% of livestock, but 9% of game species as lynx and caracals, prey on the smaller size classes of annually (Graham et al. 2005). Where wild ungulates livestock such as sheep and goats, while the largest are utilized by people for either commercial gain (com- felids prey on the full size range of livestock. Interme- mercial hunting and game farming) or for leisure diate-sized felids, such as pumas and leopards prey on (sport hunting), competition and conflict may occur smaller livestock and juveniles of the larger species. between users and large felids. For example, cheetahs The spatial distribution of livestock depredation is in Namibia kill economically valuable wild ungulates often uneven. Stahl etal. (2001a, b) found that in the on commercial game farms, leading to persecution of French Jura Mountains, there were consistent 'hot the cats by farm owners (Marker era/., Chapter 15, this spots' of lynx predation on sheep. The areas were volume). Similarly, hunters in the Russian Far East often also areas with high roe deer (Capreolus capreo- come into conflict with tigers over real or perceived lus) densities, suggesting that lynx populations may competition for ungulates (Miquelle et al. 1999a). have been sustained by reservoirs of natural prey. European lynx are persecuted when they compete Similarly, in southern and central Brazil, livestock with sport hunters for prized ungulate trophy species were most vulnerable to puma and jaguar predation (Breitenmoser etal., Chapter 23, this volume). within or in close proximity to forest fragments or riverine forest, habitats that provide felids with cover and residual levels of natural prey (Mazzoli et al. 2002; Palmeira et al. 2008). In southern Africa, live- Livestock depredation by felids stock predation by lions and leopards is often higher Felids are obligate carnivores. Humans and felids come close to the boundaries of protected areas (Fig. 6.2). into conflict in ecosystems, where a high proportion of This was found to be the case adjacent to Khutse ungulate biomass is made up of domesticated species. Game Reserve, Botswana (Schiess-Meier et al. 2007) For the most part, the smaller felid species do not cause and Chirisa Safari Area, Zimbabwe (Butler 2000). In >, CO c s= =5 CD 3 CO CD CO o •a c c CD an CO tlfl c CO CO CD o C c g CD CO CO C CO CO 'w E =5 'o 3 3 *± C c o .2 E to CD CO CO O -I CO CD CO If g. 3 ro ^1 CD CD .a c o E CD =5 CO >, cb a. CL CD CO C 0) =5 g CD CD CD 3 3 E 'co CD O CD ^f o CO o o 3 CO to CO CO o CT CD co •a c O CO tXO CO m -a o >1 o CO CD co CD CO CO -Q Q. CO O _%: o ^c CO o o O Q o Q_ o | CO J= =5 JD o CD O to CD CD CO o CD CD CD o O •a "a CD CO V Q CL o c O c ^~ CO "CD o CO CO CO "co O 3 o CO g E •as 3 o •a "co CO c CD CD CO c CD o CO •on CO o •o CD CO CO "co -C CO o o O 'co CD CD o =5 CO CD .C Q. CO o =5 CO Q. CO CO >, Q. a> UO O CD CO c O CD O =5 "CD CO CO o CD CD CD Q co O T3 CO E _i e Q. a. < CO o. -a

CO CO CD o CD CO a. >> O Z> CD CO 'co CO CO CO CO CD" S Q. o e o CO ^ o CO 'CD a. o o" =3 O 'co CO CD "co ID O E H •a o O c o o CO C o O E CO CO CO •a O o Z> Z) CD to O CD 6? 4- CO CO a. c CO 3 3 CD 3 CD o Q. =5 c CO o o •a CO CN o 00 o o o c > CD <* 3 jD J= 6b CO CO CD o CO c CO CO CD o c o % CO CD CD % CL CO CO O CD" c CD o CD o CD CO CO JZ O o CD o CO =5 « 8 CO CO CO CD to 6? &9 to 'TD •a c •C CD ro g LO CM to o o CD O ^ to CO CN 00 •OB CO J= CO CO •ao •a c o CD c o o co CO o o c o CO vp ^ — gP CN CO co > g cK C CD to O = CO CO CO =5 o CcO 03 -sf CO e o ^f CO I CD O Q. y 9 o £. 00 CD T3 CD O If CO o c ^ CO o d ^ ri -8 * CN d ro co O O O o ID co >CD •a a) oo i£ co a> -D °0 - T3 0> CO LO 0) CM c = 00 jg 00 •a % a> _a o &^ u CO 0) . CO • ro ro __ CO co O ,n CO CO CD CO "D CD -^ t. <-- « (5 OsS-= 13 u 1 co co O CO CN o ° ••-' ro ^o «' rom coro +r -t- -t- o ° o CO CD 3 ^ a^SccOCoS-SgCDco CD E co CUB ro P0 ro S- > CO o c - LO __ a o CD ^ _i "a oo ro = ro o ro 00 U J ri co s m co o CD 5 ro ^ OJO« sco o co o CD CO co to o CD CD C3 d d a. - C CD CC CD d O ro O d d -D a.

"D C CD O E ro "S ro -a o CO . ro C c S. % O CD § o E jB o § o 1 Zi 0) ro u Zi ^> ro o CD — 13 O 3 t- o to 0) o CN 05 o 4- o CD CM o I I CO CD o CD O) CO o CD O) o to o CD en o E 1 Q CN CN CC CD X j_, Q. w ro oo" 3 co O o o 0) ro ro CO „ CO c ro O d CO o (C o c CO c LU O) - CD •*• CD LO g ro CD ^ CD c o JS: O CD CD 'Q. CO CO _C ^ 3 o o CO CD to CO CD 4= K^ 8 > Z r >, CO CO CD CO C " Si,CD CD c c CD CD CO CO CC CO o o 3 CM CO T3 CO S- E y _l CD ^ 5 H CD h- ^ CC a •a JS: co c o •a •a CD =5 CD tXO SZ o Q. g g CD 5P CD o CO c a. o CD 3 f c ± CD c o CD CD SZ CM "O _ . .C CD a) SZ E ttf) (3) J E CDm 3 o 3 CD S= X Ofl "O CO 12 O £> CD co CD CD HI 111 y £> o o 10 d _ c O -a "- h- 3 tuO o J= CO CD ^ ^g CD c 's= co c 2 CO to co CM •£ CO 3 — .g o m =5 w "* CM 2 in -a -Q co o vH •§" CO CD o CD CD -O c i o o c/) •a "O CO LO 3 3 CL JS: si 0) o o CD 3 o B CL 5 g CO LO co C CD E j*: CD =5 CO CO E CO o o O o o o o CD g 0) -w *" '^ o o CO o CD _x: 1 f to CD d) CD c "CD 0) C/) to CO .2 co CO o CO N — T3 m T3 5% c c CO c T3 co co CO CD CD CD Q CD t o CD CO O CD > 01 O) LO CO CO V —1 Q_ T3 •H vH CL O a. 01 a) < Jl_i D. o i z co •a •a CD -> OJO CD co" CO CO CO CD CD ^ Q. CO O g o g 2 "a £ o CD CD •- a) o CD o sz 8 te I CO CD o CD 3 P CD CO S CD O tuO CD =5 1 _co O CD •QD - m •H ^ CO SZ I 01 > CM -- <" 9 O O \ N 00 CO ^H CD CD 3 •sT CM \ CO CD —' O co to CO vH — E CM O 5 t 00 LO i; O » CO i; .2 vH o oo c/> ,8 .2s =)« tem 3;s

CD CD CD w CO o CD T3 c/) O c CD •M CD CO CO 4-< o CD 5s "a CO CD i- CO ro CD CD cn CD -7-5 «) £0

o d a) co c 2 CM ^ -D T3 T3 CO co m tl 2 o V c o co c CO CD CD •a o -o O ci o CD o 1 CO CD CO CD 4^ sp /A -t- CD CD CD "O 5= g O CO co CD CD CD CD o CD .2 a) T3 a. Hi«? CL Q. 00 in •a CD M o tXO CM CD H co oi I 2 CD- m = LO -a CM vH "D S 2 to _ h- CD CD CD = d ^ CO CO -D 2- -^ o aT > 4= co 2 CD CD CD -a H sz CO O) d CD O rl +i >> W "2 t! vco co C ° CO CO CD c/) CD CO CD CD - 3 o oo cu ID •a c <2 -b" "a to ° o CO &1 CD CD 00 CO = CD 2 O "O CD H CD o .2 CD = P c co ^ 8 co .c .92 <- .s. -o o _l ^ X. -=- CD

EL o CO Q CO d O CL o d d o d co

•a C JZ T3 "D C CD CD C CD CO CO CL "CD S= C O CD O CD o JZ o o o

in in in CO ? i ? T en CM CM CO oo O 01 01 01 O 01 01 CM

g 4-1 BO CD CD CD CD CD CD CD 3 c CO _ "CD £ co c z - 3 ^ -* JD 3 CO CD W CD CD CD 2 f % m CO E .r- 5, C CO OJO O CD c c ^ CO C CD C E ?' CD *± O i#| .5 Oo_ £ 5 c? _^ a. ^ k O Z) O > CD > M CD S .2 en T3 c CD en CD o a> en o I to CL c T3 en c CD o CD oo c tj O 5 3 CO a> o en CM o CD en o o CD to EL o h- $ 00 o o CO en A .Q c CD o o CO CL txo en CD g Q_ > O e •o =5 "co o CL CD co ro en o en CL CO •a o o o S o CO o CD "T CD to O c CJ a. CD •a O CD CD c CJ en CO en 3 s| , CO CD O > d 00 CJ 'o5 CO c _J o 00 o en o o OO CD CD >- O 5, 0) _g oo CD CO en •a CD CD CD 5 CL CD 03 QO ^ 2 •a o -a 00 a. O O CD CO LO CD r < c CD •an "CD 00 U} CO E JZ C o en CL OJ "S -E o CO 'JZ "co CD S CD o C 'CD c CD JZ 3 4- a. 5 "o CD CL CD o T3 £< Q. CD CD o CL S= CD CD "CD o o .£= a F CD CO CO > js: CO £ I s "co so CD 'co CO rs CL CD CJ c o O CN CO c 89 CO o in CJ CD CD CJ c a. C CO &9 _>, CO ^T •H CO w O a. ^f CD •o LO E o "CD E m E O ^f 3 JZ CD C CO o 00 a "O %£ CJ CJ CD CL C iff CD CO c CO •y oo" o CO 3 CM OB n o m o CO O ^1 O CD c JZ tj E CO E en 00 o o CD a. CD o CD CO CJ CO LO CD CJ > CD CO O 3= c o LU o a. CJ U_ CO ^H CM CL txo o c a) C/) 3 W o CD O CO CO O 00 3 C CD CD g en o CJ > o o _CD o "co "co en OO CO CD to _C 3 5? CD SZ CD OD CO O H CO en o ir to -a CO to T3

J^ o .5 o CD cr £ co o o CO ^ i O 00 CD o ;„ CD IT) to JZ CO CD _a o CD •o CD > a> > en to <; CD "co OO OO 00 jH O "CD e c " CJ z o o CO o ~w JZ o c c 2R ° CD CD CO -Q en en •o 3 •^a .^o CD to o Q_ -a -£= o 3 co to CD CJ cq o CD c c CD 9 (D o -a CM _g a. CO d CO CM 0)

CO 3 4- "O CO 2 en to en O CD CO CD en o 2 CO • - "D ^ CO \ o o CD f CO CD CD c *i M- .oo E CO J >, CD "co O0 - CD .AI CD CD r\T- CO CD o JZ a> CO CD LO cii J5 •a ^ co en E O T3 en CD 9 E 3 CD CD > 3 $ ^ O _ CO CD CD > CD CO c _c CO O 0) O O vH £= o CD O CO s o Q_ ^ CM CN 6 CO sz >, OJO

00 CO > o en o 0) CD o CO o CO en CO CD CD o o > Q. CJ O. o >-" C5

•a § T> ^ CD 5 CD i_ CD S CD c o 9P o co a> C CD co »

•a c co CO CO CD CD -a 01 O) o CD 00 CD I I I I =5 a> o CO LO CM M- 00 c o 00 CD 00 "CD CD CD c o CJ) CD o Q CN ^1 ^1 o o —. CD ^ c CO CN g co T3 co en CO H s. c "CD •a •a (C co c C CD E O CD 3 CO c o 3 £ 3 co o £ -%: —" CL CD tj o co q co % LO CD CD en c -a O- c s= op -5 ro CD c CN OO CD% O) CD C o CO £c > CO CD 00 J= z < CJ to T3 CO CD c 0) o to CO o s CO CO co ° O I - CDS OJ1 _co o — CN CO g co -o c A O co co E m > g « LO •a CN .« tXO — O 01 CD CO -D o I .i= -a o a) $ 0, CO q= 00 c .2 CO o "= M« S co CO c 2 .9 CDtt c — « o O CO 00 o W II 00 CD g 5 % I n cJE JO 0> a. -o to co CO o a. _> X3 C t- CO o 2 CD O CD g » t! CD CD <*• CL CO = Q. t m oo cu CO OJO CN . Q- •* CO "- eg co ^ c I -a 4-, O CD CO .C CO c c £ ao If) CD CO 5= CO CL E T3 Q. vH -S CO CO co -c O § 0) CD 3 00 = P 8 O p o 2 X CO Q. a. "a •H _^: CO Q.

CN P0 CN P0 ^^ -o <* If) X CD CO « r to o — o Of a^ CD O) to o > 00 > Gl 5^5 "co CO CD O 3 co e o w o w CD CD on m CO .>> CL CD co c to CD T3 c E c" CD o CO tuO T3 CO ##{ c CO co .c 3 CD c CO o co co CD m CD CO -a CO c T3 E <£ CD to CN CD 00 CD a CO o E CN -£Z CN CN 00

00 "D op to 4- CO CN o CO S S) d 1-i o i Q. C I to CD O CO 9- 'ta i; co CN to p o oo d to o o o 01 o £«co to Q- O a? CN •a CD a) CD CD C) c c to > co o ti -^ ,, co co .a i!o Q. CO o. 0 -o co CN CD ^5 d. CD CO to CO CD CO 13 3 01 CD ^ M op 2 ao o = 2 CO vH CL a. co OO ^ CL

EL d CO CO d o o no ii

sz •a CO s= o 8 CD CO CL CO Q. CO CO o E 3 =5 CO c ro EL —i CO a.

• SS CN O o CO 4- O 01 01 o CN S I o 4 CO 00 o o CO 01 o Ol OJ o 01 01 o CO CN o to $ o a) SE 3 CO s= o CO CO CO CO c E < - " o _L C O S. CD o CD 2 o o O c SZ o £= =5 o QJ 00 5 % % n C to N 0

losses to lions and leopards were lower during drought years when wild prey were in poor condition and there- fore easier for predators to capture (Ogada era/. 2003). Peaks in livestock predation may also be due to seasonal availability or vulnerability of livestock. In areas with marked seasonal variation in climate, live- stock may be in poor condition during the lean season, which may make them easier prey for felids during this period (Hoogesteijn etal. 1993; Butler 2000). Neo- nates of domesticated species are often more suscepti- ble to predation than adults. This vulnerability can result in increased levels of depredation during the lambing or calving period. Peak losses to jaguars and pumas on cattle ranches in southern and central Brazil and central Venezuela often coincide with the calving season (Polisar et al. 2003; Michalski et al. 2006a; Palmeira etal. 2008). Seasonal changes in husbandry or herding practices may also result in periods of vulnerability for livestock. In France, peak losses (May-November) of sheep to lynx coincided with periods when sheep were kept in fenced fields for the entire 24-hour period, presumably increasing their availability to lynx and therefore the probability of depredation (Stahl etal. 2001b). Villages around Bhadra Tiger Reserve, India, experienced lower levels of livestock loss during the harvest season, when Figure 6.2 A village headman displays the remains of a it is thought that increased human activity in sur- cow killed by lions, Tshlotsho Communal Land, adjacent to rounding fields may also inadvertently provide protec- Hwange National Park. © AJ. Loveridge. tion for stock (Madhusudan 2003).

northern Cameroon, levels of lion depredation were inversely correlated with distance to Waza National Economic impact of livestock depredation Park (Van Bommel etal. 2007). Impacts of felid depredation on livestock vary, de- Temporal patterns of livestock depredation are often pending on the scale of livestock ownership, husban- predictable and in many cases vary with levels of avail- dry techniques, livestock type, stocking density, and able natural prey. In many areas of Africa, livestock density of predators. Small-scale, subsistence livestock depredation peaks during the season when natural owners are often disproportionately impacted by prey is most difficult for predators to acquire. This is losses, in part because they may lack resources to pro- usually the wet or rainy season, when prey are in better vide effective protection for their stock. Loss of even condition and often more widely dispersed (Patterson an individual animal to small-scale livestock owners et al. 2004; Bauer and Iongh 2005; Kolowski and Hole- has a proportionally higher impact on herds or flocks camp 2006). Hemson (2003) found that in the Maka- than the same loss to owners with more stock. For this gadigadi ecosystem, Botswana, livestock-raiding lions reason, Ikeda (2004) found that small-scale yak (Bos killed fewer livestock during periods when migratory grunniens) herders in Kanchenjunga Conservation wildebeest (Connochaetes taurinus) and zebra (Equus Area, Nepal, were more heavily impacted by snow burchelli) were present in their ranges (Loveridge et al., leopard (Panthera undo) predation on their herds Chapter 11, this volume). In Laikipia, Kenya, livestock than wealthier, medium- to large-scale owners. People and wild felids: conservation of cats and management of conflicts 171

Snow leopard predation in the Spiti Region, India, 1.7% was lost to disease, malnutrition, and misman- amounted to losses of between 25% and 52% of agement (Palmeira et al. 2008). In contrast, around annual per capita income for small-scale farmers Bhadra Tiger Reserve losses of stock to predators were (Oil et al. 1994; Mishra 1997). In Africa, subsistence around 4 times higher than losses to disease or starva- farmers in Kenya and Zimbabwe lost between 11% tion (Madhusudan 2003). In Gokwe, Zimbabwe, levels and 12% of annual income to lion and leopard of loss of livestock to predators (~5%) were similar to depredation (Butler 2000; Ogada era/. 2003). Villages losses to starvation and disease (Butler 2000). on Koyake Group Ranch adjacent to the Maasai Mara Reserve, Kenya, lost approximately US$1890 to lions and US$984 to leopard depredation over 14 months Attitudes to livestock depredation (Kolowski and Holecamp 2006). Tiger depredation People's attitudes to and tolerance of depredations by on livestock around Bhadra Tiger Reserve, cost vill- wild felids vary widely. Antipathy towards carnivores agers 16% of annual income (Madhusudan 2003). may be a result of historical or cultural attitudes, as well Around Jigme Singye Wangchuck National Park, as based on past experiences and personal values. Al- Bhutan, tigers and leopards caused losses of 17% of though generalizations are difficult across widely vari- annual per capita income, however for households able socio-cultural circumstances and it is often difficult actually affected by losses this represented up to 84% to distinguish between the underlying reasons for neg- of annual cash income (Wang and Macdonald 2006; ative perceptions people hold towards predators, there Wang 2008). These losses are significant impacts on is some evidence that levels of tolerance livestock own- the livelihoods of people, who often have few alter- ers have for predators are related to magnitude and native means of earning a living. impact of losses. In the Pantanal, Brazil, tolerance of In contrast, large ranches lose relatively less to depre- jaguars by ranchers was at least partially explained by dation. Large ranches in Alta Floresta, Brazil, lost rela- levels of livestock loss (Zimmermann et al. 2005a). tively fewer cattle to depredation than intermediate- However, other studies indicate that personal and soci- sized ranches, with no ranch losing more than 1.24% etal beliefs and perceptions may be equally important of its herd (Michalski et al. 2006a). On large-scale in shaping responses to conflict with predators (see ranches in Kenya, it is estimated that the cost of main- Cavalcanti et al., Chapter 17, this volume; Murphy taining a lion was US$290 in southern Kenya and US and Macdonald, Chapter 20, this volume). $360 in Laikipia (Patterson et al. 2004; Frank et al. Economic circumstance may also dictate levels of 2005), while the cost of maintaining a leopard cost tolerance. Subsistence farmers in Annapurna Conser- ranchers US$200-211/year in Laikipia (Mizutani vation Area, Nepal, who suffered significant losses to 1997; Frank et al. 2005). The cost of tolerating occa- depredation held highly negative attitudes towards sional depredation was considered to be low by these snow leopards and considered total extermination of relatively wealthy ranchers (Mizutani 1999). Losses these felids the only solution to the problem (Oil due to depredation by carnivores are often much smal- etal. 1994). However, in the Indian Trans-Himalaya, ler than losses due to disease, poor husbandry, or theft. where people had alternative incomes and lower de- On a Laikipia ranch, average loss of stock to predators pendence on livestock, they were more tolerant of over a 23-year period was 34 cattle and 66 sheep a year. snow leopards (Bagchi and Mishra 2006). Subsis- However, annual losses to disease were 2.5 times tence pastoralists on group ranches in Laikipia, higher for cattle and nearly 4 times higher for sheep. Kenya, were more likely to attempt to eradicate pre- Losses attributed to theft were similar to losses to car- dators than neighbouring commercial ranchers nivores (Mizutani 1997). In Kweneng district, Bots- (Woodroffe and Frank 2005). In this case, commer- wana, 0.34% of livestock was lost to carnivores each cial ranchers tolerate losses to large felids both be- year (although losses to individual owners were higher cause of a cultural appreciation of wildlife and at 1.0-11% of holdings). However losses to disease, because wild felids have value to ecotourism opera- starvation, and accident accounted for 2.8-12.6% of tions (Mizutani 1997, 1999). In other areas, people stock (Schiess-Meier et al. 2007). On central Brazilian are willing to tolerate moderate losses. In India, vil- ranches, 0.4% of stock was lost to predators, while lagers often graze livestock on government forest 172 Biology and Conservation of Wild Felids lands and are willing to tolerate moderate loss as the accidents for instance. However, because of the terri- price of access to grazing resources (Srivastav 1997; ble emotional consequences attached to the loss of Karanth and Gopal 2005). human life, human killing puts conservationists in a difficult moral dilemma. It makes conservation and protection of big cats difficult for conservationists to justify to local people who suffer losses. Baldus (2004, Human injury and fatality caused by felids 2006) gives an example of a man-eating incident in Perhaps the most striking relationship people have the Mkongo Division, Rufiji district, adjacent to Sel- with felids is that large felids occasionally prey upon, ous Game Reserve in Tanzania, that illustrates the kill, or injure people. Evidence that hominids have traumatic disruption to people's lives. In this case, at always suffered depredation by felids comes from the least 34 people were killed and 10 injured by lions Swartkrans Caves in South Africa, where the exca- over a period of 20 months, with most incidents vated skull of a hominid child bears the tooth marks occurring in crop-lands south of the Rufiji River, of a leopard (Brain 1969; de Ruiter and Berger 2000). which divides the district. Eventually, almost the en- Carnivores from the families Felidae, Canidae, and tire human population had fled to the northern Ursidae are all on record as killing people (Quigley banks of the river to escape the lions, leaving homes and Herrero 2005). Due to the need to overpower and crops unguarded. Leaving aside the emotional and kill their victims, human-killing felids are invari- and psychological consequences of living alongside ably the largest species, with lions and tigers being man-eating carnivores, the perceptions and attitudes the most dangerous and widely publicized (Table of people towards wildlife and conservation of large 6.2). Famous historical cases of man-eating big cats carnivores are seriously impacted. In poor countries, include the man-eating lions of Tsavo, which, in compensation for loss of human life is often grossly 1898, are reputed to have killed around 128 con- inadequate. In Tanzania, compensation for loss of struction workers on a railway line being built from human life to wild animals is around USS30-50 (Bal- Mombasa to Nairobi, Kenya (Patterson et al. 2003). dus 2004). Apparent undervaluation of human life Other historical examples include man-eating tigers may further compound ambivalent attitudes of local and leopards in northern India, one of which (a communities towards conservationists and conserva- tigress) reputedly killed 436 people before being tion managers, reduce public acceptance of big cat hunted and shot (Corbett 1946). Boomgaard (2001) conservation, and undermine conservation efforts. reports that in India, in 1875 alone, tigers and leo- Indiscriminate retaliatory killing of cats often results pards killed 1039 people. However, human deaths from incidences of human killing. In the case of the due to large cats are not a thing of the past. Human Rufiji maneater, wire snares were set throughout the deaths and injuries caused by lions are relatively area to catch the culprit, killing at least eight lions common in central and southern Tanzania, with (which may or may not have been maneaters) and around 563 Tanzanians killed between 1990 and probably many other non-target species, before the 2004 (Packer et al. 2005b). Similar numbers of people actual maneater was eventually shot (Baldus 2006). are killed and injured in the Sundarbans of India and Attempts to eliminate maneaters may impact the Bangladesh, with 294 people killed between 1984 viability of local populations of carnivores. and 2001 in the Indian Sundarbans (Siddiqi and Choudhury 1987; Karanth and Gopal 2005) and 79 people killed between 2002 and 2007 in the Bangla- The magnitude of the problem desh Sundarbans (Khan 2007). Table 6.2 provides some examples of accidental killing, injury, or predation on people by felids. Felids from the genus Panthera are the most consistent maneaters, Human attitudes to man-eating and their although this behaviour is rarely recorded in P. onca consequences (Rabinowitz 2005). Pumas (Puma concolor) are known At a global scale human killing by felids is relatively to attack and kill people, but incidences are extremely rare, much rarer than human deaths due to traffic rare, with 10 deaths and 48 injuries throughout the CD 3 CD CO •a T3 •g c CO o Q. •a O O eo ro CD CD CD 0) o 5 = CN 3 3 oo E 2 -^ l J= J£ en •a 3 O c oo en o en I E op CO 3 eo CD c g|s CD O i- O •C CD CD c -C to o - to E a> C = O) CD CD ^ OO > o eo CD CD ^2 ^ § a. %3 3 O LJ CD c n ao o ro CD 2 T3 0) o C CO en CO — ft CD J= Sp. •a g Q. -O CD O eo 5 O CD & ro g CO ° 2 CO CD CD Q. T3 CD CD = -1 0) eo c en — S CD -° en a. .*: J= CD en CD CD "D >, S c^ CD o T3 E oo CD a> *- •a CD CO CD 03 -° 2? CD _: 3 M= -O o CO E oo ro CD O ^ o *± 3 O g CD o c 2 o — Q. E =3 en ' ft — ro O •a o c * CD •a CD B i CN - c c CD « on CD .Q 3 =5 00 N E o CO O •a O CD ^ 0) CD co •" g — S g o CD CD s= 0) CD CD ^3 o CD ro c ro Cfl a. E on c o CD p to CD § s > m co o o. CD C a> CD CO en >, g CO T3 = -o -a on fl CD > 03 CD o C CD C c = JZ ^ CD •a = CD cz en °° CD CD -S y CD _ o = CD CD =5 en "a -c If) CD C c CD CD ro oo o o -cY .5 i_ 3 •a CD ® o J= OO t Q. o CD CDS2 ^> C CD ro o "o a. CD ro 3 O CD o. O CD J= ^ O O Q. en = c 11 a o en o C CO ! S ^ Q. CD — CD CD 3 ro tn ° -* CD CO CD = 0) O O CO « co <. "O — o OO o en CD -^ >, CO g . .e 2 as CD c o c s= = C = g Q- CO ro as <" CD CD co -o in OO CD CD o 2 2 CD -e O ±± jn •^ ft <° ro on CD 3 § § CN ID « C CD "O CD CD a. O CD CD ro g 2 Q- -a "O f CO CD a. S 13 Q. C 3 a. •v§ CD o a> E = c CD O CD o CO CD m -worn 3 O 3 = CD -C CN 3 E c c ro Q < O "5t OO LO Q- 3 £ B « 00 co Q. O O ( -a s= a Q Q Q a Q CD M CD < < < < < < -Q E =5 CD * S= ttf) < E CO JZ 3 ^ _ o u O LO 05 CN O CO CD O 05 CO 4- t! O iri CD LO o 1 d d 1 < vH LO CM in 1 00 H 1 Cfl C C CO CD JZ o E % o CD CO 00 CD 3 CD o CN 4" in CO o CN CN 00 M oo x -a rl 4" CN oo LO CN 00 ^H CN CO

CD .CD 00 LO 3 O N CJ) CO LO X CD H CO 11 c & = to CD -c -a E en oo c g 3 3 o Q. CD 3 E o X 00 _i C CD g "o en c '•*-' t; CD c c c c c c CD O CO CD CD CO CO CD ro ro CD o o o o o "a 4-; CD c c c .5# c ao 2 "5 Q. o O H— O 4- O O g g CO o CD Q. r CO 3 O CD "a u ^ C CD tLO CD CD CD SZ > co" CO c tXO _CD 03 O C =5 > •a CO o CO O =5 o C CD 3 O CT> CD •a O CD ro > CL 1 03 to O CO 03 CD CD o O C c "CD CL O CD C S= CD 3 _c £ CD •a h- 00 CD o > CD "CD CD •a E UO •a CO CD 03 CT> E % C3 -a •a CD co ^H c CD CD 03 CD E o 3 o JD •2, o t 3 •a tLO 3 CL CD o 03 o SZ 03 o? s= - -9 Q. CD T3 E CD T3 O jb CO CD M- ^^ •OB CD c Q. c o ajCD rl CO > CD CO CD T3 O CD •a = 00 CO 03 T3 CD CD 03 CD •a 03 o CD 00 03 03 c ^ co =5 •oh CD .5P CD 3 CO to -a "O CD O CD O o c CL CO O 03 c CD CO SZ CD tXO o CD CL H CO CD e LO •a o o o 3 CD c CO s= co CD 3 CO CD •a CO c CD o CD CO CD "TO CD H o CD o £ CL 03 c to CD 03 CJ o 3 E as CD O O CD CD 03 03 CD CD o CD o

a* Q Q Q I— CD < < < Q < <

CD 2 CO

o o CM o 4" d LO CO CN d

CD J= 4" I % 10 o ^ CN O) LO 03 00 4- x -a CO H CN CN LO 00

CD .0) c E <-- 03 CO LO o > o CD CN LO Z UO CO

co c 00 CD CN "O CN LO co CD o a.CD CD CD O rr CD CD CD CD 03 CD CD Q. 0) 3 Ofl 00 Ofl UO Ofl tLO no CO

LO o O O o CN a> O o O o CO CO o CD CN CN CN 00 o CN 03 en I I I I I CD O o LO in CO CN o oo 03 05 00 CO LO O 00 "CD 01 00 O) O) O) O) o 03 cn T3 Q •H •H CN •H CD 3 C o O o r- LO C Is CD CD c O CD , LO in .£2 E c O J= CO 39 "D CD CD CD I CO N CO c CO c CD % C c CD CO CD 3 CO 03 (0 T3 "D -Q -a f O §1 CD* CD « > i? CD J5 CD CD O # CO O CD ao "D -- oo -a "CD .C CO CL C C CM c c CL s s 3 'ID CD D £ CD 3 CD 3 m C CD -k CD CC N iir lihi m co ^ CO CO CO People and wild felids: conservation of cats and management of conflicts 175

United States and Canada from 1890 to 1990 (Beier counters between people or livestock and felids are 1991). Lions and tigers, the largest living cats, are res- more frequent because either felids disperse from ex- ponsible for most incidents, with episodes of leopards isting or emerging populations or when people enter as maneaters being occasionally reported, particularly felid habitat. Conflict may also be more likely to occur in northern India (Corbett 1946; Athreya 2006; when levels of natural prey have been depleted. These Athreya et al. 2008). Levels of human killing by felids three situations are discussed in more detail below. vary widely. Human fatalities to tigers were 0.01 peo- ple/1000 km2/year in the Russian Far East, where human population density was low and interactions Dispersal of felids into marginal, with tigers rare and usually accidental (Miquelle et al. human-dominated habitat 2005a). This contrasts strongly with the 50 people/ Ironically, the consequence of successful protection 1000 km2/year killed by lions specifically targeting of populations of large felids is that surplus indivi- people as prey items in Rufiji district (Baldus 2004, duals may need to disperse out of habitats or refugia 2006). By their nature, incidents of human killing by already occupied by conspecifics. If the surrounding felids tend to be clumped in space and time, so compil- areas provide no connectivity to other habitat ations of incidents across large spatial scales or long patches and/or are marginally suitable habitat, al- time periods tend to under-represent the local impact. ready used by people, conflict with human popula- Likewise, reports that include incidents over a short tions in these areas can result. time-span and localized scale may overstate the global Examples of this include tigers dispersing into importance of man-eating as a behavioural trait of buffer areas surrounding Chitwan National Park, felids, while at the same time capturing the traumatic Nepal. This has led to 88 human deaths between impacts such incidents have on human communities 1980 and 2005, with many of the dispersing tigers living alongside populations of large felids. being old individuals displaced from their ranges It is not always possible to unambiguously distin- (McDougal 1993; Gurung et al. 2006a). A similar guish between those incidents that were the result of example of competition for space leading to displace- accidental encounters between a person and large ment into areas of human settlement occurred in felid and those where the person was viewed as a Lupande Game Management Area, Zambia, where a prey item by the felid in question. Karanth and pride of lions, displaced from South Luangwa Na- Gopal (2005) compare the examples of Kahna Tiger tional Park by a stronger pride, killed three people Reserve, India, where human deaths are relatively (Yamazaki and Bwalya 1999). Conflicts between peo- low (but not infrequent), with the Sundarban Tiger ple and large felids over livestock loss are common Reserve, India, where human killing by tigers is fre- on the boundary of many protected areas in Africa quent. In Kahna, tigers occasionally come into con- and Asia. Large felids disperse into marginal bound- tact with people in the densely populated areas ary areas and kill livestock. For example, dispersing surrounding the park, leading to occasional human lions kill livestock around Etosha and Hwange Na- deaths when tigers are disturbed or accidentally en- tional Parks (Stander 2005b; A.J. Loveridge, personal counter people harvesting forest products. However observation). Similar conflicts between lions, people, in the Sundarban Tiger Reserve it is clear, both from and livestock occur in the boundary areas of Gir the high incidence of human deaths and the circum- Forest, India (Saberwal et al. 1994). Problems of dis- stances of the attacks, that people are the intended persing carnivores from emerging populations have prey of some tigers (Table 6.2). been experienced in western Europe, where reintro- duced European lynx have come into conflict with sheep farmers (Kaczensky 1999). Characterization of human-felid conflict Situations when livestock depredation or man-eating Human encroachment on felid habitat are more likely to occur fall into a number of broadly People and their livestock enter (legally and illegally) defined circumstances. These are scenarios when en- large felid habitat for a number of reasons, the most 176 Biology and Conservation of Wild Felids common being for hunting or extraction of forest may increase the chances of people encountering products, use of grazing resources, and occasionally pumas and account for the increases in puma attacks for other reasons such as leisure activities, tourism, or on people in the past decade-and-a-half (Cougar as refugees (Johnson era/. 2001b; Kruuk 2002). These Management Guidelines Working Group 2005). Peo- activities bring people into contact with felids and ple also become vulnerable when circumstance may result in either accidental killing of people (and/ forces them to intrude into big cat habitat, as illu- or felids) or provide opportunities for felids to prey strated by the example of Mozambican refugees, flee- upon people or livestock. In the Sundarbans of Ban- ing into South Africa, via the Kruger National Park, gladesh, there are no permanent settlements in the being killed and eaten by lions (Kruuk 2002). forest but people use the forest for extraction of for- est products both legally and illegally. Here some tigers apparently regularly prey upon people, with Prey depletion causes felids to switch fluctuations in numbers of fatalities correlated with to alternative prey periods of high use of the forest (Khan 2007). Simi- Evidence from some studies suggests that felids have a larly, injury or death caused by tigers in the Russian preference for natural prey. This may be because avail- Far East appears to occur when people accidentally ability of domesticated species is not reflected by their encounter tigers while hunting in the forest (Kerley numerical abundance, as livestock is often protected etal. 2002; Miquelle etal. 2005a). Livestock losses are and livestock-raiding animals risk higher levels of mor- high in areas where livestock are grazed in or in close tality in human-dominated environments. Eurasian proximity to forested habitats with resident felid lynx in Norway did not select habitat patches with populations (Wang and Macdonald 2006; Palmeira high livestock densities, but preferred areas with high etal. 2008). In Norway, sheep are grazed unprotected roe deer density, and preyed predominantly on roe in forested habitats, leading to high losses to carni- deer, despite the fact that sheep densities were eight vores (Swenson and Andren 2005). times higher (Moa et al. 2006; Odden et al. 2006). Another reason people encroach on felid habitat is Rabinowitz and Nottingham (1986) found that to settle in these areas. Nyhus and Tilson (2004b) jaguars prefer natural prey to domestic stock. Hemson record the increasing incidence of livestock loss and (2003) found that lions appeared to 'prefer' wild prey, man-eating by tigers in Sumatra during the 1970s- at least during periods of high wild ungulate abun- 80s when settlement of the island by transmigrants dance. In this case, they killed livestock less than ex- was encouraged and people encroached on tiger hab- pected based on availability in the good season and in itat during the massive transformation of the low- proportion to availability in lean season. Accordingly, land rainforests into oil palm plantations. However, it was suggested that maintaining suitable levels of incidents declined during the mid-1980s as tiger natural prey in protected-area buffer zones may serve habitat was destroyed and tiger populations on the to limit depredation on livestock. island declined. This pattern of conflict followed by A common suggestion in reports of depredation population decline has been closely documented for on both livestock and people is that levels of natural tigers on the Indonesian islands of Java and Sumatra prey have been depleted and this may provide an and in India between 1600 and 1950 by Boomgaard explanation for these behaviours. While this has (2001). Similar patterns of depredation by tigers may rarely been quantified, a number of examples appear have occurred across south-east Asia as human pop- to support the suggestion and may explain a propor- ulations increased and colonized forest habitats tion of the cases. For example, Asiatic lions attacked throughout the nineteenth century (McDougal people around Gir Wildlife Sanctuary, when natural 1993). Rapid fragmentation and conversion of for- prey densities declined after a prolonged drought ested habitats to agricultural land in Amazonia bring (Saberwal et al. 1990; Saberwal et al. 1994). In the people and felids into contact and result in high case of man-eating lions in central and southern levels of conflict (Michalski et al. 2006a). Similarly, Tanzania, their natural prey was depleted, forcing the increasing encroachment of human residential the lions to switch to prey (bush pigs, Potamochoerus development into puma habitat in North America porcus) found in proximity to human settlement. People and wild felids: conservation of cats and management of conflicts 177

This may increase the likelihood of encounters with them, leading to higher rates of predation. However, people guarding their crops, potentially leading to there are few data to directly support this suggestion. man-eating events (Packer et al. 2005b). Hoogesteijn Higher rates of livestock killing by male felids could (2002) and Crawshaw (2004) found that loss of nat- also be due to larger male home ranges and wider ural habitat and prey predisposes jaguars to kill live- dispersal distances and therefore an increased likeli- stock. Similarly, losses of domestic stock to snow hood of encountering livestock (Linnell et al. 2001). leopards were high, making up around 58% of This is certainly the case in lions, a species where snow leopard diet, in areas with low abundance of subadult males often disperse widely. Patterson et al. natural prey (Mishra 1997; Bagchi and Mishra 2006) (2004) found that 66% of stock-raiding lions in south- ern Kenya were subadult males, while Funston (2001a) records that of 100 livestock-raiding lions in ranch land adjacent to Kgalagadi Transfrontier National Mitigation of conflict Park, 59 were subadult males. In this case, a further 31% of raiders were females with cubs, suggesting the Characteristics of problem animals possibility that increased nutritional requirements of Defining a large cat as a problem requiring a solution lactating females or females feeding dependent young depends on both where it is and how it is behaving. may predispose them to stock raiding. Linnell era/. (1999) distinguish between two kinds of In a study on the boundary of Etosha National Park, 'problem animal'. The first are animals that are in the Namibia, Stander (1990) was able to distinguish be- 'wrong place' (generally in a wide-scale matrix of tween habitual and occasional stock-raiding lions. habitats, where not all individuals have access to Around 70% of habitual raiders were adult males, livestock within their home ranges). Individuals dis- while 54% of occasional raiders were subadult indivi- persing out of protected areas or individuals coming duals. He found that 11 of 12 'occasional' raiders trans- into contact with livestock or human intruders into located did not reoffend; however, 'habitual' raiders their habitat might fall into this category. The second quickly returned to their home ranges and livestock 'type' of problem animal is one that kills more live- raiding unless they were translocated > 100 km. stock (or people) per encounter than do conspecifics. Infirmity and disability have often been cited as a Animals with old injuries or habitual livestock raiders possible reason for man-eating and livestock depreda- (Stander 1990) could be included in this definition. tion and there are a compelling number of examples of There are a number of factors that appear to pre- maneaters and livestock raiders with old injuries or dispose individual felids to becoming problem ani- damaged teeth. This may or may not cause the animal mals. Male felids appear to be more likely to become difficulty in capturing or subduing wild prey. For ex- stock raiders (Stander 1990; Cunningham era/. 2001; ample, over 50% of 17 man-eating tigers from Chit- Funston 2001a; Odden et al. 2002). Odden et al. wan had old injuries caused by gunshot wounds, (2002) found that male Eurasian lynx were not only intraspecific fights, or had worn or broken teeth (Gur- more likely to kill livestock than females, they were ung etal. 2006a). One of the two 'man-eating lions of also more likely to make multiple kills during a raid- Tsavo' had deformities of skull and jaw (Peterhans and ing incident. Specific male behaviours may also be Gnoske 2001) and the maneater of Rufiji was a 3.5- factors that influence whether or not livestock is year-old male with severe abcessation in its lower jaw taken (Linnell et al. 1999). Bunnefeld et al. (2006) (Baldus 2006). A number of the notorious man-eating found male lynx were more likely to be found in tigers and leopards shot by Corbett were old and had proximity to human habitation than females with been suffering from disability (Corbett 1946). In post- kittens, which may explain both higher male mor- mortem examinations of Belize jaguars, Rabinowitz tality and predisposition to livestock killing by males (1986) found that of 13 livestock raiders, 10 had old of this species. Similarly, because felids are markedly injuries (many caused by old gunshot wounds), while sexually dimorphic, with males being larger than in a sample of 17 non-livestock killers all were healthy. females, one might expect that male felids would Hoogesteijn etal. (1993) report similar trends in Vene- have a greater size range of livestock available to zuelan jaguars, where 10 of 19 cattle killers suffered 178 Biology and Conservation of Wild Felids from old gunshot injuries. The notion that old age or icy provides a set of protocols that clearly delineate disability could be the cause of aberrant predation on appropriate actions for the various situations likely humans or livestock is appealing. While this appears to be encountered and guide the process of determin- to be the case in some incidences, it is equally likely ing the appropriate response options. While every that old or infirm individuals lose competitive intra- situation is unique, a general protocol that empow- specific interactions in prime habitats and are thus ers local wildlife managers to make decisions, makes displaced from core areas and into marginal habitat, them accountable, and provides response teams with where they come into contact with people and live- guidelines to follow for choosing the appropriate stock. Habitual livestock raiders may have been more course of action is a critical first step in dealing quick- likely to have been shot at in the past. Indeed, Patter- ly and efficiently with problem animals. son et al. (2003) found that the skulls of lions killed as Because local support for, or at least tolerance of, problem animals around Tsavo National Park, Kenya, large felids is one of the key factors determining the were no more likely to show signs of disability, or fate of all wild populations, elimination of real or tooth breakage than those in museums, which were perceived threats is pivotal. The ability to retain a presumed to have been collected at random. problem animal within the wild population will de- pend on the abilities of the response personnel, ef- fective liaison with local communities, and the Solving and mitigating human-fetid conflicts severity of the problem, as well as whether official Designing conservation policy and building management interventions are adequately directed, resourced, and capacity supported. In developed countries where there is a strong com- mitment to carnivore conservation, compensation schemes have been established to mitigate losses, Lethal control coupled with selective removal of individual prob- Predator reduction or elimination, either through lem predators. In many developing countries, lack of state-sponsored predator control or unregulated kill- attention by governments or low and delayed com- ing, has historically been the method of choice for pensation discourages livestock owners from partici- protecting livestock. In this way, large felid popula- pating in similar schemes where these exist. The only tions have been extirpated from large areas of North perceived solution is to kill the predator, and often and South America, Africa, Europe, and Asia. While many non-target animals are killed in the process. complete eradication of predators is no longer prac- People who fear for their safety or perceive that they tised in many areas, reduction of predator numbers are at economic risk will not support conservation or limitation of population growth through lethal efforts. control methods may be compatible with manage- Conservation practitioners need to design frame- ment plans that include zonation of predator man- works for implementing a comprehensive response agement. to problem animals. Such frameworks often include Where complete eradication of predators is not creation of professional problem animal response desirable, lethal control of specific problem animals teams and crafting of national policies and protocols can be used to deal with livestock raiding or inci- for response to conflicts between humans and large dents of human death or injury. However, it is some- felids. Response teams should be professional, well times difficult to identify the individual animal trained and adequately equipped, and have the ex- responsible and indiscriminate lethal control may perience and confidence to handle the range of situa- kill many non-target individuals or species, particu- tions that are encountered and fully understand the larly if wire snares or poison are used. Control meth- consequences of their actions. Many countries have ods that specifically target problem animals, such as no written policies for dealing with problem carni- toxic collars on vulnerable livestock (Burns et al. vores; situations are handled on a case-by-case basis 1996), use of dogs or skilled trackers to follow prob- with no guidelines and without the proper staff lem animals, and shooting or trapping of culprits training, equipment, and resources. A national pol- when they return to recently made kills may provide People and wild felids: conservation of cats and management of conflicts 179 more focused control of problem individuals (Odden lem animals appears to be a limited and temporary et al. 2002; Woodroffe and Frank 2005). solution (Stahl et al. 2001a; Herfindal et al. 2005a). Trophy hunting is sometimes advocated as a Furthermore, identification of the problem animal means to reduce predator densities or to target spe- after the fact is not always straightforward and in cific problem animals. Trophy hunting has been situations where tourist hunting opportunities are shown to temporarily reduce population densities sold commercially, bogus claims of problem animals in some felid species (e.g. puma, Lindzey et al. need to be guarded against. Lethal control of specific 1992; Eurasian lynx, Herfindal et al. 2005a; Stoner problem animals may be better undertaken by pro- et al. 2006) depending on the sustained level of the fessional problem animal control personnel, rather hunting pressure. than the sport-hunting public; the former is likely to Felid populations often recover quickly from pop- be a more targeted and timely response. This is par- ulation reductions, if there are nearby source popula- ticularly the case when dealing with man-eating pre- tions (e.g. lions, Smuts 1978; Eurasian lynx, Stahl dators (Cougar Management Guidelines Working et al. [2001a]). Therefore, unless hunting is used on a Group 2005). consistent basis to limit population growth, the ben- Where problem animals are part of a rare or efits to livestock owners are likely to be temporary. endangered population, lethal control may not be Furthermore, vacant home ranges are often filled by the option favoured by conservationists. Neverthe- dispersing subadults, a demographic group that ap- less, particularly in cases where people's lives and pear more likely to become problem animals. There livelihoods are at risk, removal of a problem animal is evidence that ranges made vacant may be divided by lethal control may be the most expedient and between multiple subadult dispersers resulting (at efficient option available. Unsuccessful or ill-con- least temporarily) in increased population density ceived interventions or unwillingness on the part of (Laing and Lindzey 1993). This may escalate rather managers or conservationists to deal effectively with than resolve conflicts. This being the case, sustainable problem animals may lead to a perception that the sport hunting which seeks to maintain viable predator welfare of animals is valued over the lives and liveli- populations (and therefore hunting opportunities) hoods of local people. In this case, support for con- may be incompatible with predator reduction (or servation efforts may be undermined (Tilson and eradication) in livestock raising areas, unless livestock Nyhus 1998). owners are prepared to tolerate some losses. In an alternative approach to reducing population Translocation density, Anderson (1981) describes proactive culling Other options for removal of problem animals in- of potential livestock-killing lions in Hluhluwe-Um- clude translocation back into protected areas, to folozi National Park, South Africa. It was found that zoos or other protected sites. This intervention has animals dispersing from the Park and causing live- been used in a number of areas, particularly with stock losses on surrounding farms were predomi- lions and leopards in Africa, but success is equivocal nantly subadult males (18-42 months old, 59 of 79 due to high post-release mortality, extensive move- problem lions destroyed). Proactive culling of sub- ments, and homing behaviour of translocated ani- adult males in the Park resulted in a greatly reduced mals. In Tsumkwe district, eastern Nambia, Slander incidence of livestock loss on surrounding farmland. et al. (1997a) translocated six livestock-raiding leo- However, highly targeted interventions such as this pards a total of 12 times to sites 10-135 km from may only be practical in intensively managed situa- their ranges. All individuals returned within 2 days tions, such as small-fenced reserves where animals and killed livestock again in an average of 8.2 (range are often regularly monitored and individually 1-20) months. Man-eating leopards translocated recognized. from Pune district, India, to other forests and re- Reducing felid population densities through hunt- serves continued to reoffend at the new sites ing can provide temporary benefits to livestock own- (Athreya 2006) and similar behaviour by leopards is ers through reduced depredation. However, use of described in east Africa (Hamilton 1976). Of 14 hunting as a tool to selectively target specific prob- stock-raiding male lions (two territorial adults, two 180 Biology and Conservation of Wild Felids non-territorial adults, and 10 subadults) translocated possible in developing countries. Although translo- distances of 50-85 km from their home ranges in the cation of problem felids is in general unsuccessful as Kgalagadi Transfrontier Park, South Africa, all except an intervention, livestock-killing cheetahs (Acinonyx three subadults returned and killed livestock again jubatus) and problem Amur tigers have been success- within 170 ± 37 days. Most were repeatedly translo- fully translocated. In Zimbabwe, in the mid-1990s, 21 cated (between two and six times) but either re- cheetahs were moved from the south of the country turned or were destroyed while killing livestock ~450 km to Matusadona National Park in the north, (Funston 2001a). Similarly, 40% of the 54 lions trans- and successfully established a population there (Pur- located from around Etosha National Park returned chase 1998; Purchase and Vhurumuku 2005). Of four to their home ranges (Stander 2005b) and for the Amur tigers captured and translocated after killing same reason capture and repatriation of stock-raid- domestic livestock or attacking people in the Russian ing Asiatic lions failed to control livestock raiding Far East (Fig. 6.3), two were successful in that they around Gir Wildlife Sanctuary, India (Saberwal et al. caused no further conflict with people, killed wild 1990). Similar post-release behaviour has been ob- prey, and survived their first winter (Goodrich and served in other large felids. Rabinowitz (1986) re- Miquelle 2005). ports that two livestock-killing jaguars translocated As well as translocating problem animals to alter- to Cockscomb Basin National Park, Belize, soon left native wild habitats, some problem animals are oc- the park. One continued to kill livestock and was casionally moved into captive settings. Five of 37 shot after 5 weeks; the other disappeared. An experi- man-eating tigers from Chitwan were captured and mental translocation of 14 cougars in New Mexico housed in zoos (Gurung et al. 2006a), but facilities to resulted in 25% mortality within 3 months and, house large numbers of such animals are limited and overall, nine animals died during the study period. quickly become saturated (e.g. for tigers in Nepal, Two mature males moved back ~477 km to their India, Malaysia, and Indonesia; leopards in India original ranges in 166 and 469 days. Eight cougars and Sri Lanka; and pumas in North America). Fur- moved >80 km back towards their original range thermore, wild-caught felids do not generally settle before establishing new home ranges (Ruth et al. well in captivity (Karanth and Gopal 2005). How- 1998). ever, movement into captive settings might be justi- Translocations are most likely to be successful if fied for highly endangered species, especially if problem animals are moved long distances across release of offspring into the wild could enhance rein- significant landscape barriers to areas with reasonable troduction programmes at a later date. prey densities and few livestock or people. However, In addition to limited success, translocation opera- such areas are infrequently available and tend to have tions are costly and require high levels of technical already established populations of the species being expertise and logistical support. Funston (2002) translocated. Translocation of non-territorial, sub- estimates that repatriation of lions that broke out of adult individuals, which would disperse anyway in a Kgalagadi Transfrontier Park cost in excess of US$218 natural setting, appears to be the most likely to be per lion. Karanth and Gopal (2005) note that tigers successful. In pumas, translocation was most effec- are often injured or disabled during translocation, tive for animals aged 12-24 months and least effec- often through damaging their teeth in steel cages tive for older individuals (Ruth etal. 1998). Based on that are not adequately designed (Fig. 6.4). On the results of lion translocation operations in Nami- balance, translocation does not appear either partic- bia, Stander (1990) recommended that habitual prob- ularly successful or a viable tool for management, lem animals be destroyed while occasional raiders, particularly in countries with limited resources. particularly subadults, could be rehabilitated by translocation. While in this case translocation activ- Protection of livestock and improved livestock husbandry ities reduced the number of lions that had to be Historically, traditional livestock husbandry prac- destroyed, it required extensive record-keeping and tices have sought to limit the availability of livestock expertise to capture, mark, and re-identify stock rai- to predators. However, in areas where predators have ders. This kind of intensive management is rarely been extirpated this knowledge has often been lost. People and wild felids: conservation of cats and management of conflicts 181

Figure 6.3 An Amur tiger is released back into the wild by WCS and Inspection Tiger after being rescued from a poacher's snare. © J. Goodrich, Wildlife Conservation Society.

This is the case over much of western Europe, where that were free-roaming (Mazzoli et al. 2002) and traditional herding methods such as herd guarding by around Jigme Singye Wangchuck National Park, Bhu- shepherds and guard dogs and use of secure enclosures tan, livestock losses were high in villages that did not have been largely abandoned. This has contributed to have stables or corrals, or where livestock was grazed in conflicts between farmers and re-emerging predator forest habitats (Wang and Macdonald 2006). populations such as Eurasian lynx (Kaczensky 1999; Protective bomas (corrals; Fig. 6.5) and herd guard- Stahl et al. 2001b). Where livestock are grazed exten- ing (often by young men and boys) are widely used sively without supervision or protection, losses to pre- across Africa to reduce livestock losses (Frank et al. dators are liable to be high. For example, in Norway 2005). Schiess-Meier et al. (2007) found that over a 3- sheep graze unsupervised in forested habitats, where year period only three of 2272 livestock-killing inci- they suffer high levels of predation to lynx and other dents (largely by lions and leopards) took place inside carnivores. In contrast, sheep are usually kept in bomas in central Botswana. Similarly, Ogada et al. fenced enclosures in Sweden and loss to predators is (2003) found that livestock losses were reduced with limited (Odden et al. 2002; Swenson and Andren attentive herding, strongly built bomas, and the pres- 2005). Similarly, free-roaming cattle in central and ence of guard dogs and people. Enclosures constructed Amazonian Brazil and central Venezuela were subject of poles or wicker were superior to those built of to relatively high losses to jaguars and pumas (Polisar thorny Acacia branches or wire mesh, because they etal. 2003; Michalski etal. 2006a; Palmeira etal. 2008). reduced the chances of stock panicking at the sight By contrast, in southern Brazil, managed flocks and of a predator and breaking out of the protective enclo- herds suffered lower levels of depredation than those sure. Similar reductions in depredation resulted from 182 Biology and Conservation of Wild Felids

Figure 6.4 An African leopard trapped in a steel cage trap. Care needs to be taken in design of such traps to avoid damage to the felid's teeth and claws. In general, trapping and translocation of large felids is not a successful or viable management intervention. © A.J. Loveridge. stone and wire mesh corrals built to protect sheep Improved animal husbandry may also reduce live- against snow leopards in Ladakh, India (Jackson et al. stock losses to predators. Free-roaming cattle in areas 2002). However, Kolowski and Holecamp (2006) such as the Pantanal, Brazil, and Llanos of Venezuela found that on ranches adjacent to the Masai Mara are often in poor condition, increasing their vulnera- Reserve, Kenya, leopards preferentially targeted stock bility to jaguar predation (Quigley and Crawshaw in enclosures, particularly those built of poles, but that 1992; Rabinowitz 2005). Vulnerable stock such as in general fences, guard dogs, and human activity calves and sheep often require more intensive man- deterred predators. Guard dogs can be used to protect agement. Peak losses to livestock owners often occur small livestock (Linnell etal. 1996). In Nambia, farm- during the calving or lambing seasons (Polisar et al. ers using livestock-guarding dogs reported a 73% re- 2003; Michalski et al. 2006a; Palmeira et al. 2008). In duction in livestock loss to cheetahs (Marker et al. areas of South America where puma and jaguar preda- 2005a). Similarly, llamas {Lama glama) and domestic tion is a problem, use of pastures away from forest buffalo (Bubalus bubalis) have been used to protect fragments and close to human habitation is recom- flocks and herds from pumas and jaguars (Crawshaw mended for calving enclosures (Quigley and Craw- 2004; Hoogesteijn and Hoogesteijn 2008). Visual and shaw 1992; Palmeira et al. 2008). In addition, acoustic repellents, such as bright lights (Sechele and management of breeding in cattle herds aimed at lim- Nzehengwa 2002) and noises (e.g. shotgun blasts, iting birthing to a short period may allow for intensive playbacks of bio-acoustic sounds such as sounds of protection for part of the year and also act to swamp barking dogs; Koehler et al. 1990), may be useful in predation (Crawshaw 2004; Palmeira et al. 2008). In deterring carnivores. However, carnivores habituate areas of Norway where sheep are vulnerable to lynx quickly to unusual stimuli and it is not clear whether predation, Linnell etal. (1996) recommend shifting to such methods are effective in the long term. Deter- cattle farming, as cattle are not preyed upon by lynx. rents such as electronic shock collars and taste aver- Losses in productivity often far exceed losses to sion work reasonably well under experimental predators. Cattle raised under semi-wild conditions conditions, but have not been widely applied to man- have extremely low productivity, with pregnancy age livestock raiding (Frank and Woodroffe 2002). rates reaching only 40-50% and have very high People and wild felids: conservation of cats and management of conflicts 183

Figure 6.5 Livestock kept at night in a secure pole 'bomas' or corrals suffer lower levels of depredation than those left out to graze. © J.E. Hunt.

neonatal mortality and mortality to disease (Hooges- without the requirement to demonstrate adoption of teijn 2002). Improved husbandry not only limits op- approved preventative measures (as is required in portunities for predation by felids, but also reduces Sweden; Swenson and Andren 2005) can lead to a losses to disease, accident, and theft and therefore perverse incentive to neglect livestock. There may also the potential for losses due to poor management also be incentives to wrongly attribute losses due to to be blamed on felid depredation (Hoogesteijn poor husbandry, accident, or disease to carnivores in 2002). order to claim compensation. Furthermore, compen- sation may act as an agricultural subsidy, providing Compensation incentives to expand farming activities with the po- Damage compensation schemes provide one ap- tential for conversion of additional natural habitat proach to mitigate damage caused by carnivores. (Nyhus etal. 2005). Long time delays between occur- Payment of compensation for damages has the effect rence of damage and payment of compensation and of spreading the economic burden and financial risk bureaucratic difficulties in making claims may also between those who must live alongside carnivores be problematic. Saberwal (1990) found that 81% of and those who wish to see wildlife protected. Com- villagers suffering losses to lions around Gir Wildlife pensation for actual carnivore depredation (some- Sanctuary did not bother to make claims, due to times called post-damage compensation) is widely procedural bureaucracy. Similarly, failure to fully used in Norway, Sweden (Swenson and Andren compensate for losses may hamper the effectiveness 2005), and India (Karanth and Gopal 2005). How- of compensation schemes. For instance, around Bha- ever, while it is clear that compensation may allevi- dra Tiger Reserve, where significant losses of live- ate some of the costs and potentially promote stock to tigers occurred, low levels of compensation tolerance of damage caused by predators, this ap- were received (3% of estimated losses, with only 20% proach suffers from some major disadvantages. Verifi- of claims successful; Madhusudan 2003). Corrupt or cation of damage (to eliminate fraudulent claims and poorly managed compensation schemes may deepen over-estimates of loss) can be time-consuming and antipathies and mistrust of managers and conserva- expensive and lead to animosity between conserva- tion efforts. Furthermore, to provide a sustainable tion managers and livestock owners. Compensation long-term solution significant financial resources 184 Biology and Conservation of Wild Felids are required (Miquelle et al. 2005a). For instance, in torted in that there is no disincentive to protect Norway, compensation costs 5 million Euros/year livestock from predators. Similarly, because payment (Linnell and Breseth 2003). Schemes are rarely feasi- is made on the basis of expected damage, there is no ble in countries with limited conservation resources time-lag in payments, which eliminates uncertainty unless supported by international sponsorship or and promotes transparency and trust. Nevertheless, non-governmental organization (NGO) funding. transaction costs remain high, as carnivore popula- Unfortunately, few evaluations of post-damage com- tions still need to be independently assessed and pensation schemes have been undertaken (Nyhus et agreement needs to be reached over levels of pay- al. 2003; Nyhus et al. 2005). It is not known if they ment. In common with post-damage compensation are cost-effective and there is little evidence that payments, viability of schemes requires significant compensation payments are effective in improving funds over extended periods. Care needs to be people's tolerance or reducing attempts to eradicate taken that payments do not encourage new immi- predators. In one of the few analyses available, Haz- gration into the area or subsidize increases or expan- zah (2006) found that compensation did not im- sion in livestock ownership. prove attitudes to lions or prevent retaliatory killing on Mbirikani Group Ranch, Kenya. Alternative livelihoods, benefit sharing, and stakeholder Private insurance schemes to cover costs of carni- participation vore damage have also been attempted; however, in Antipathy towards carnivores may be a result of his- many cases rural farmers are unwilling to cover the torical and cultural attitudes as well as based on ex- relatively expensive premiums required (Nyhus et al. periences of loss or damage. These attitudes may be 2005). This unwillingness may stem in part from difficult to change or ameliorate. However, there are concepts of ownership of wildlife and the perception examples where stakeholder attitudes to both conser- that compensation is the responsibility of the gov- vation and presence of carnivores have been im- ernment or wildlife management institution. How- proved by access to alternative revenue or livelihood ever, success of insurance-based schemes has been choices, benefit sharing, and local participation. claimed in improving tolerance of snow leopards in Co-management and stakeholder participation Pakistan, although this scheme was subsidized appear to improve attitudes towards conservation through ecotourism revenues (Hussain 2003b). Via- efforts. In reindeer husbandry areas of Sweden, car- bility of insurance compensation schemes may rely nivore surveys jointly undertaken by reindeer own- on innovative financial mechanisms such as this. ers and conservation managers reduced conflicts An alternative to post-damage compensation is over estimates and encouraged transparency and co- conservation performance payments or compensa- operation (Swenson and Andren 2005). Participation tion in advance (Schwerdtner and Gruber 2007; of local communities in prioritization, conservation Zabel and Holm-Muller 2008). Such payments are planning, and implementation enhanced local pro- made based on the expectation that damage is likely tection of snow leopards as well as improving the to occur and is usually linked to successful conserva- feasibility and sense of ownership of livestock pro- tion outcomes (e.g. reduced levels of carnivore mor- tection initiatives in Hemis National Park, India tality and protection of carnivore habitat). In the (Jackson and Wangchuk 2004). Likewise, inclusion Sami reindeer (Rangifer tarandus) husbandry area of of local landowners in conservation meetings in the Sweden, conservation performance payments are Pantanal, Brazil, improved perceptions of jaguar con- made on the basis of the number of certified repro- servation efforts and engendered ownership of pro- ductions of lynx and wolverines (Gulo gulo). The cesses and decision-making (Rabinowitz 2005). level of payment is calculated from the expected Oil et al. (1994) found that villagers who benefited losses due to each carnivore during its lifetime and from ecotourism were more willing to tolerate the amount to around US$29,000 per individual off- presence of snow leopards. There is a higher toler- spring (Zabel and Holm-Muller 2008). The advantage ance of this species in areas where people are less is that payments are made regardless of damage; dependent on livestock and have access to alterna- therefore, livestock protection efforts are not dis- tive livelihoods (Mishra et al. 2003; Bagchi and People and wild felids: conservation of cats and management of conflicts 185

Mishra 2006). In Namibia, ranchers who raise live- communities, which may undermine the effective- stock using ecologically sound management techni- ness of initiatives (Walpole and Thouless 2005). ques gained access to niche markets and premium Furthermore, revenues earned through ecotourism prices for 'predator friendly beef. Benefits from in- may be enthusiastically accepted in the short term, itiatives such as this may increase tolerance for but in the long term may be invested in develop- cheetahs and other predators on ranch land (Marker ment or expansion of agriculture (Murombedzi and Dickman 2005; Marker et al., Chapter 15, this 1999). The key here is that benefits from ecotourism volume). On commercial ranches in Laikipia that schemes need to be clearly linked to the need for benefit from ecotourism or have wealthy foreign in- sustainable conservation of ecosystems and natural vestors, landowners are more tolerant of depredation resources. on livestock than are ranches that gain nothing from tourism. Similarly, traditional pastoralists do not tol- Zonation of land use erate predators at all: lions dispersing into these areas Not all landowners or communities will tolerate pre- from commercial ranches are promptly poisoned dators, and certain livestock management practices (L. Frank, unpublished data). However, residents of and human-dominated or urban landscapes are in- communal areas that hope to profit from future wild- compatible with the presence of predatory felids. life tourism profess greater tolerance than those who Thus geographic differentiation of land-use areas, are not planning conservation-based enterprises ranging from complete protection, through areas (Romanach era/. 2007). Stander etal. (1997a) describe where predators are tolerated and population man- an innovative ecotourism venture that combined use agement and/or utilization occurs to areas where of traditional skills, revenue generation, and ecotour- predators are not tolerated, seems a sensible way of ism. In the Kaudom area of Namibia, local people focusing conservation efforts and resources, while experienced losses of livestock to leopards. A pilot at the same time recognizing the importance of ecotourism venture took advantage of the local Ju/ people's livelihoods. Hoansi people's exceptional tracking and bush-craft The goals of zoning conservation of carnivores are skills. They were able to track and find leopards for to conserve viable populations of predators and to paying tourists in 92.9% of attempts. The two villages minimize, or at least mitigate, conflicts with people engaged in leopard-tracking tourism earned N$ 10,005 (Linnell et al. 2005). Limiting the interface between (~US$2140) in a year. This revenue amounted to people and large carnivores can serve to reduce the 12 times the value of livestock lost to leopard over areas where conflict occurs. This allows prioritization the entire district. of mitigation efforts and efficient use of conservation However, in many cases claims that revenue from resources in these areas. Enforced zoning schemes ecotourism or tourist hunting alleviate conflict and can be used to prohibit certain human activities, improve attitudes towards conservation are almost though forms of resource use or extraction that do entirely untested (Walpole and Thouless 2005). In- not result in habitat modification such as fishing, centive schemes are often heavily subsidized by ex- regulated logging, regulated harvest of ungulates, ternal bodies (Mishra et al. 2003; Miquelle et al. leisure activities, and tourism may be compatible 2005a) and internalizing costs to ensure sustainabil- with conservation of felids (particularly those that ity is often challenging with uncertain outcomes. do not pose a threat to human life). In some cases, Wildlife revenues appear to work well in incentiviz- the presence of charismatic felids may increase the ing protection of habitat when landowners are rea- desirability of an area as a leisure or tourist destina- sonably wealthy and where owners have land tenure tion. The conservation of smaller species may be and control over resource access and use (Bond et al. compatible with land uses that exclude the presence 2004). However, there is less incentive where resource of larger predators. For instance, small felids pose no ownership is centrally or communally controlled, threat to livestock and coexistence may be possible if land tenure is less secure, or where livelihood alterna- no habitat modification occurs. tives are absent. Ecotourism revenues can be misman- Many southern and east African protected area net- aged, misappropriated, or subverted by elites within works have been designed to incorporate areas of strict 186 Biology and Conservation of Wild Felids protection (national parks or reserves), and areas be managed are designated as 'source' or 'sink' popu- where wildlife can be sustainably utilized (often lations. Sport-hunting opportunities are provided in through trophy hunting). In addition, community- the sink areas, while source areas are protected from managed areas (known as game management or wild- utilization. Management using a landscape of life management areas) have been established in some sources and sinks to provide both adequate protec- countries (e.g. Botswana, Zambia, Zimbabwe, and Na- tion for the species and opportunity for utilization mibia), where wildlife is tolerated and utilized for the may prove to be an effective strategy for conserving benefit of communities (Lewis and Alpert 1997; wide-ranging predators. However, understanding of Hulme and Murphree 2001). Utilization areas are a species' behavioural ecology is crucial. Predators, often set up as buffer zones between national parks particularly large felids, are wide ranging and mini- and agricultural areas. Zonation in other regions is mum protected-area sizes need to take this into ac- often less clear-cut. For instance, many areas of Europe count if edge effects are not to have negative impacts are already inhabited by people. In these areas, initia- on population viability (Woodroffe and Ginsburg tives to prioritize conservation of predators have not 1998; Loveridge etal., Chapter 11, this volume). always met with success and have in some instances been abandoned (Linnell etal. 2005). Establishment of reserves without adequate separ- ation (e.g. with fences or natural barriers such as Exploitation of felid populations large rivers) of people and wildlife has led to intense through trade and utilization conflict on the peripheries. Clearly, management of Human cultures have always prized products derived conflict in areas where predators and people are ex- from large carnivores. Traditional cultures and fash- pected to coexist makes zonation of predator conser- ionable society value felid furs to advertize personal vation a challenge. In areas where people and status and trade in furs has had significant impacts wildlife share the same landscape, it would appear on felid populations. Felid products are used to 'cure' that coexistence is most likely to be successful under illness, to ward off misadventure, and to bring good circumstances where disadvantages due to conflicts . For instance, tiger bone as an ingredient of are outweighed by benefits of tolerating wildlife. Traditional Asian Medicines (TAM) is thought to Some conservation practitioners have advocated cure rheumatism, weakness, and paralysis (Mills separation of people and predators as the most viable and Jackson 1994). In south-western Nigeria, serval solution, particularly in areas where human densities (Leptailurus serval) flesh and tongue are believed to are high. Nyhus and Tilson (2004b) reported that Way- cure leprosy and rheumatism, while leopard skin is Kambas Reserve in Sumatra successfully protected used to treat snake bites (Sodeinde and Soewa 1999). tigers and eliminated conflicts with the surrounding There are recent reports about a growing illegal mar- human communities. This was largely because the ket for tiger meat as an exotic cuisine (Damania et al. reserve excluded people and reserve borders are largely 2008). Utilization of felids through trophy or sport formed by rivers that created boundaries seldom trans- hunting can provide motivations for conserving gressed by either tigers or people. Karanth and Mad- habitats and wildlife populations. We discuss the husudan (1997) and Karanth and Gopal (2005) exploitation of felid populations through trophy support the voluntary relocation of people from the hunting, fur trade, and for traditional medicines, vicinity of tiger reserves to reduce conflicts and protect and the potential threats and impacts these have tiger populations. Such voluntary relocations to make on conservation of these populations. space for expanding tiger populations have met with success in the Terai Arc Landscape, Nepal (Seiden- sticker etal., Chapter 12, this volume). In the western United States, zonation provides the framework for management of puma popula- Trophy hunting tions (Laundre and Clark 2003; Stoner et al. 2006). Hunting large felids for sport has occurred for Here, subpopulations within the meta-population to thousands of years, with records of lion hunts going People and wild felids: conservation of cats and management of conflicts 187 back to the time of Pharaoh Amenhoteb III in 1400 taken belief that removal of carnivores would 'pro- BC and medieval tapestries depicting European no- tect' ungulate populations, thereby providing bles hunting lions and other felids (Guggisberg improved opportunities for hunting. Along with 1962). Indian and Nepalese royalty hunted exten- loss of habitat, decline in prey populations and pred- sively and records show the astounding number of ator eradication initiatives, trophy hunting may tigers they killed for sport. The Maharajah of Surgu- have contributed to historical declines in some jah is reputed to have shot 1150 tigers in his lifetime, large felid populations. Indeed, I.R. Pocock (1939: followed closely by the Mararaj of Gwalior and his p. 219) wrote: 'In all parts of the world occupied by guests, who accounted for 900. British civil servants Europeans where lions occur, the disappearance of and military officers on leave hunted frequently, lions is merely a question of time.' with some individuals credited with killing hundreds Historically, hunting of carnivores was often char- of tigers. The British Royal Family did their bit when acterized by lack of any controls or regulation (at least visiting India and Nepal, with King George V killing for the elite) and in some senses was indistinguish- 39 tigers in 11 days in 1911 and the Prince of Wales able, in motivation and effect, from predator eradica- shooting seven over a 4-day period in 1922 (San- tion activities that were prevalent at the time. khala 1978; Mountford 1981). During the 1960s, Contemporary trophy hunting (also known as demand for tiger-hunting trophies accelerated as sport, tourist, or recreational hunting) tends to be tiger populations dwindled and as it became appar- more restrained and harvests are often strictly regu- ent that hunting tigers would become officially pro- lated where competent management authorities are hibited (Sunquist and Sunquist 2002). in place. In addition, trophy hunters are often In colonial Africa, carnivores were viewed as ver- concerned about and promote conservation values min and killed wherever possible (Fig. 6.6); this (Lindsey etal. 2006). Felids are popular quarry species hunting was, at least partially, motivated by 'the where trophy hunting is legal, and large charismatic thrill of the chase' particularly among the colonial species attract large trophy fees (usually collected by elite hunting in the more remote areas of east and government or management authorities). Trophy southern Africa. Early hunting expeditions to the hunters pay premium prices to hunting guides or Serengeti typically killed large numbers (sometimes operators to hunt large felids. For example, puma hundreds) of lions (Turner 1987), often in the mis- hunts in New Mexico cost US$2000-3000 per hunt

Figure 6.6 Early settlers in east and southern Africa eliminated many of the larger predators from the land they colonized. (Photograph courtesy of National Archives of Zimbabwe.) 188 Biology and Conservation of Wild Felids

(Logan et al. 2004) and before hunting was officially for trophy-hunting activities. For instance, in addi- banned, hunters paid US$11,200 to hunt a snow tion to national parks and protected areas, 1.4 mil- leopard in Mongolia (Anonymous 1989). In Bots- lion km2 are used and, to varying degrees, protected wana, in 2006, tourist hunters paid US$140,000 to for trophy hunting in sub-Saharan Africa (Lindsey hunt a male lion, with US$100,000 of that going to et al. 2007). Crawshaw (2004) argues that allowing the community in the district it was killed (J. Rann, trophy hunting of problem jaguars may provide in- personal communication), although lion hunting centives for landowners to manage habitats to en- has since been banned in Botswana (Fig. 6.7). courage self-sustaining populations of this species. Revenues generated from trophy hunting can However, trophy hunting can have demographic benefit local communities (if explicitly channelled) consequences for felid populations. Adult males are and provide employment for specialists that provide often targeted and high male turnover can result in hunting-related services (e.g. hunting guides and high levels of infanticide, which may in turn reduce taxidermists). This can provide justification and sup- reproductive success within the population. If severe, port for conservation of populations of felids and this can lead to population decline (Greene et al. 1998; other species at local and national scales. Revenues Whitman etal. 2004). Gross (2008) found that heavy can also be used to manage populations, protect hunting of male pumas in Washington, DC, United habitats, or to compensate livestock owners for States, led to severe perturbation including few resi- losses, thereby improving levels of tolerance and dent adult males, influx of immigrants from surround- acceptance of predators (Loveridge era/. 2007b). Fur- ing home ranges (leading to widely varying puma thermore, habitats are often set aside and protected densities), increased infanticide, and more puma- human conflicts. However, felid populations are rela- tively resilient to moderate levels of harvest (e.g. 10% of adult male lions; Greene et al. 1998), recovering quickly if immigration from surrounding habitat or nearby populations is possible. Smuts (1978) found that populations of African lions in Kruger National Park, South Africa, recovered within 17 months (large- ly through immigration) in areas that had been exper- imentally depopulated by culling of 129 lions from three experimental areas. However, the social dis- ruption caused by removals persisted beyond the 17-month experimental period (Smuts 1978). Rela- tively rapid recoveries occurred in lion population in Hwange National Park, after trophy hunting had been suspended in surrounding hunting concessions (Loveridge et al., Chapter 11, this volume). However, Lindzey et al. (1992) found that recovery of an experi- mentally manipulated puma population did not occur within 2 years when 28% of harvestable age animals were removed from the population. Trophy hunters and hunting operators (outfitters) do not always behave ethically or responsibly. There are often strong financial incentives to overexploit hunted resources, and hunting activities often occur in remote areas with little or no official oversight. For example, Spong et al. (2000a) found that in a sample Figure 6.7 The skin and skull of a trophy-hunted lion is of trophy-hunted leopards (n = 77) shot in Selous displayed in a hunter's skinning shed. © J.E. Hunt. Game Reserve, Tanzania, 28.6% were female despite People and wild felids: conservation of cats and management of conflicts 189 hunting quotas for leopards being officially restricted over 80,000 ocelot and 15,000 jaguar skins from the to males. In this case, poor supervision of hunting Brazilian Amazon in the early to mid-1960s for the activities allowed hunters to shoot females and po- fashion fur trade (Smith 1976). This massive export tentially overexploit this resource. Effective regula- of skins prompted the Brazilian government to ban tion of hunting activities to ensure compliance with export of wildcat skins in 1967. Myers (1973) estim- hunting quotas and regulations is often required to ates that the fur trade was worth around US$30 ensure that trophy hunting of felids and other spe- million in the late 1960s and early 1970s. The de- cies remains sustainable. mand for spotted cat skins was so high that, in Neo- Despite examples of overexploitation (Spong et al. tropical South America, hunting for cat skins became 2000a; Yamazaki 1996; Loveridge et al. 2007c), tro- a way of life for many local people in remote areas phy hunting appears to have relatively low impacts (Payan and Trujillo 2006). Hunting provided a lucra- on felid populations if there are large, protected tive livelihood and extraction of forest products was source populations and if management practices well organized through intermediaries and middle- and quota setting are informed and regulated by men. During this period, in Columbia, a Tigrillada carefully monitoring the hunted and source popula- (cat hunter) was typically paid around US$ 130-350 tions. Protection of habitat and the potential for for a jaguar skin, which became worth US$520 to a benefit streams for local communities and econo- middleman once it reached Bogota, before being mies might outweigh behavioural and demographic eventually sold to a New York furrier for US$2500 impacts on felid populations, although clearly these and made into a coat worth US$20,000 in a New York should not be ignored if the overall goal is to sustain Boutique (Nowell and Jackson 1996; Payan and the target populations over the long term. Trujillo 2006). The quantities of spotted cat skins extracted from South American range states from the 1960s to 1980s are staggering. In 1970, 140,000 ocelot and margay Exploitation of felids for their fur (Leopardus weidii) skins were traded in US markets, Felid furs are often beautifully patterned and humans while 84,493 oncilla (L. tigrinus) skins were traded in have, for millennia, used furs for manufacturing 1983 alone (McMahan 1986). Between 1976 and clothing, adornments, and household decoration. 1979, 341,588 Geoffrey's cat (Oncifelis geoffroyi) and The wearing of felid furs often confers status and is a 78,239 Pampas cat (O. colocolo) skins were exported symbol of the wealth and power of the owner. Masai from Buenos Aires (Mares and Ojeda 1984). Old morans (young warriors) who have killed a lion wear World spotted cats were also popular with furriers, lion mane headdresses to signify their courage. The including the leopard, snow leopard, clouded leop- fashion of wearing fur coats made of spotted cat pelts ard, tiger, and cheetah (Nowell and Jackson 1996). gained immense popularity among the wealthy of Nine thousand, one hundred and sixty-two leopard western Europe and the United States in the 1960s skins were exported from Uganda between 1924 and and 1970s. The resulting overexploitation of some i960, leading to special protection of the species in wild populations of spotted cats prompted the enact- that country in i960 (Treves and Naughton-Treves ment of the Convention on International Trade in 1999). Seventeen thousand, four hundred and nine- Endangered Species of Wild Flora and Fauna (CITES) ty leopard skins were exported to the United States in in 1975. Recently, tiger and leopard skins and 1968 and 1969 and demand for leopard skins was clothing made from skins have become popular estimated at around 50,000 skins per year in the early among the newly wealthy in China and Tibet (EIA 1970s. Furthermore, because leopards were often 2004; EIA-WPSI2006), driving an unprecedented on- hunted on an unmanaged and often illegal basis by slaught of poaching in source countries, especially tribesmen or subsistence hunters (Fig. 6.8), many India and Nepal. skins were rejected by middleman traders as being Demand for felid skins has driven widespread ex- damaged or poorly cured, implying that losses to ploitation and sometimes extirpation of felid popu- wild populations were even higher than figures lations. One famous example is the extraction of suggest (Myers 1976). 190 Biology and Conservation of Wild Felids

Figure 6.8 Demand for leopard skins led to widespread hunting of the species, with at least some of the illegal trade in skins supplied through a network of illegal trappers and hunters. (Photograph courtesy of National Archives of Zimbabwe.)

Concern over the magnitude of trade in spotted in 1973 to a high of US$164 a pelt in 1978 (Fuller etal. cats and its potential impact on wild populations 1985), with resultant harvests reducing populations of (Koford 1973; Myers 1973, 1976; Theile 2003) led this species to dangerously low numbers (Hornocker to moratoria on trade in some species (Nowell and 2008). The modern fashion fur trade relies on pelts Jackson 1996). Protective legislation was introduced from the Canadian and Eurasian lynx (I. canadensis in many range states and CITES legislation restricting and L. lynx), bobcat, and the leopard cat (Nowell and trade in many of the spotted cats, particularly the Jackson 1996). Canadian lynx furs have been larger species, was enacted in 1975. The European exploited since the 1700s (Elton and Nicholson Union (EU) banned all imports of Latin American 1942). Although trapping is currently well regulated, cats in 1986 and many of the smaller Latin American populations may be unsustainably trapped during cats were placed on CITES Appendix I between 1989 cyclic population declines (Poole 1994; Slough and and 1992. Although trade in spotted cats still exists Mowat 1996). Bobcat populations currently appear (1000 spotted cat skins were seized in Argentina in stable, suggesting that fur harvests are biologically 1990; Anonymous 1992), trade restrictions have led sustainable; however, very high volumes of leopard to a steady decline in trade (Figs. 6.9 and 6.10). For cat skins exported by China (over 200,000 skins in instance, 30,000 margay skins were exported in 1977 1987) have been cause for concern (Nowell and Jack- (most from Paraguay to western Europe) but trade in son 1996; Sunquist and Sunquist 2002). this species was reduced to 138 skins by 1985 While markets in Europe and the United States (McMahan 1986). Similarly, in 1969, at the peak of have declined due to strict legislation and negative the trade in spotted cats, it is estimated that 61,000 sentiments about use of furs in fashion items, other leopards were killed for their skins, with no more markets have opened up, particularly in a more pros- than 6000 killed by 1988 (Martin and de Meulenaer perous eastern Europe and an increasingly wealthy 1988). China. The illegal trade in tiger and Asiatic leopard However, the fur trade shifted to other species as (P. pardus fused) skins appears to have been increasing substitutes, such as the bobcat (Lynx rufus), and leop- throughout Indochina since the 1990s (Rabinowitz ard cat (Prionailurus bengalensis), with both demand 1999; EIA 2004; Nowell and Xu 2007). Poaching of and prices increasing dramatically in the late 1970s wildlife for the fur trade appears to be well organized (Fuller etal. 1985; McMahan 1986). As a result of high and lucrative. Increasing demand for skins is thought demand, prices for bobcat pelts rose from US$15 a pelt to be driven by wealthy Han Chinese who value felid People and wild felids: conservation of cats and management of conflicts 191

100,000-

10,000

o 1000

100

COCOOOJ-^CDCDOIM •*r < > (i 11 < i 11 0)0)0)0)0)0)0)0)0) m t» t» < 1 11 11 < i 11 <\l <\l <\l (\l <\l Year

Figure 6.9 Trends in international trade in skins of two species of Neotropical spotted cat (ocelot and margay). Reduction in demand and imposition of trade bans by the EU and CITES in 1986 and 1989, respectively has limited the trade. These data are likely to be incomplete, as only official exports are recorded, but, unless seized, illegal ones are not. (Source of data: CITES-UNEP.)

Figure 6.10 Jaguar skin from the Venezuelan Chaco. © A. Taber.

(particularly tiger) skins for the prestige of owning an of growing wealth among urban Tibetans. Skins are expensive and exotic item and because they are also used at traditional ceremonies such as weddings, thought to bring good luck. There also appears to and a ceremonial tent made of 108 whole tiger skins be a market in curios purchased by Western tourists was seen in 2006 at the Litang Horse Festival, in the (EIA 2004). Use of tiger and leopard skins in tradi- Chinese Province of Sichuan (EIA-WPSI 2006). tional Tibetan 'chubas' (ceremonial gowns) became However, concerns raised by environmental groups increasingly fashionable through the 1990s as a result have seen this use decline markedly (EIA-WPSI 2006; 192 Biology and Conservation of Wild Felids

Banks and Wright 2007). Symbolic burning of tiger conditions. The carcass is sold to traders who capture and leopard skin robes occurred in Tibet in 2006 fol- the bulk of the profit by smuggling tiger parts into lowing condemnation of use of wildlife products by urban centres of East Asia. All parts of the tiger can be the Dalai Lama. This may have resulted in both greatly traded with a total retail value in the region of US reduced demand and prices for these items (Huggler $10,000-70,000 (Damania etal. 2008). 2006), although Chinese officials, who discourage the In India, wildlife poaching and related crime is Dalai Lama's influence, ordered Tibetans to continue well organized, with extensive networks of suppliers to wear their traditional apparel, including clothing and buyers (Kumar and Wright 1999). The highly adorned with tiger and leopard skin. Trade in tiger and profitable nature of the trade and lack of serious leopard skins remains highly lucrative. A trader in policing and judicial disincentives in many range India is able to sell a tiger skin for US$1500 (having states do little to curtail the trade in Asian cat skins. paid a local poacher in the region of US$15), while the Although countries like China appear to be enfor- same skin sells in China for US$16,000. A consign- cing wildlife trade laws more vigorously (Nowell and ment of 31 tiger, 581 leopard, and 788 otter skins, Xu 2007), fighting wildlife crime is not a major pri- seized in Tibet in 2003 while being smuggled from ority in many South-east Asian range states. Agencies India to China, was thought to be worth an astonish- charged with environmental protection are often ing US$1.2 million (EIA 2004; EIA-WPSI2006). under-resourced and clear policies and political will Though legislation is in place to protect tigers and are often non-existent. Punishment for trading in most other Asian cats, wildlife agencies frequently lack wildlife products provides little deterrent, and fines the resources for effective enforcement of the laws. for possession of illegal skins are paltry in relation to Poaching penalties are harsh, but the likelihood of ap- the value of a single skin on the illegal market. Tra- prehension remains low and that of conviction even ders can adjust their margins to accommodate lower (Damania et al. 2008). Hunting of tigers for trade change in judicial pressure, competition, or demand is widespread and trade is thought to be more prevalent at the retail end of the market and thus frustrate now than in previous decades. This may exacerbate initiatives to diminish incentives to poach (Bulte declines due to logging, habitat loss, and expansion of and Damania 2005). Furthermore, prosecution of human populations (Rabinowitz 1999). The 783 tiger wildlife criminals is frequently delayed and rates of and 2766 leopard skins seized in India betweenl994 conviction are low. For instance, in India, which has and 2006 are thought to represent a fraction of skins on a well developed institutional structure for conserva- the illegal market, suggesting that the extent of illegal tion, only 14 convictions have been achieved in 748 poaching of felids for their skins may be extensive. EIA- cases of confiscation of wildcat skins (EIA 2004). The WPSI (2006) consider the illegal trade in tiger skins trade also spans a number of countries (Fig. 6.11), between India, Nepal, and China to be a substantial with sources in India, Indonesia, Laos, Thailand, threat that may have driven recent declines in Indian Myanmar, and Vietnam and markets on the Chinese tiger populations. Having depleted many tiger popula- tions, commercial poachers have turned to other Asian big cats: Asian lions, leopards, snow leopards, and clouded leopards. Having poached tiger populations /' i i L (0 in Cambodia, Myanmar, and Thailand, commercial Q. t India —•

Mainland, Hong Kong, Taiwan, South Korea, and of tiger bone (Hemley and Mills 1999). Domestic and Japan (Li and Wang 1999; Rabinowitz 1999; Baker international trade in tiger parts is banned in most et al. 2006; EIA-WPSI 2006). Control of this illegal countries and some countries like South Korea have trade requires regional and international commit- virtually eliminated the trade through vigorous pros- ment and transnational cooperation to tackle trans- ecution. However, the residual trade has proved boundary trade. much harder to control and eliminate. In regions where enforcement and prosecution have been in- different, tiger products are still obtainable. In a sur- vey of Yunnan Province, China, Li and Wang (1999) Use of felids in TAM found felid parts to be widely available and similar Conservationists have been increasingly concerned by surveys in Sumatra, Indonesia, found little evidence trade in wildlife products used in Traditional Asian of declining trade in tiger products, despite extensive Medicines (TAM) and remedies. Trade in tiger parts efforts to raise awareness (Shepherd and Magnus and derivatives has been banned around the world 2004; Ng and Nemora 2007). However, a recent sur- for more than a decade and law-abiding practitioners vey of 518 TAM outlets in China found that only of Traditional Chinese Medicine (TCM) now use alter- 2.5% carried tiger bone products, suggesting that natives. Yet, the illegal trade continues. The World use may be declining in Mainland China (Nowell Federation of Chinese Medicine Societies (WCMS) and Xu 2007). However, because few data are avail- has declared that tiger parts are not necessary for able from before the 1993 ban, quantification of human health care and that alternatives are plentiful, declining trade is difficult. affordable, and effective. Poachers continue to kill There are a number of captive bred populations of tigers to satisfy a stubborn demand for tiger bones to tigers, leading to the suggestion that products from make health tonics. While loss of habitat may be the 'farmed' tigers could be sold commercially to replace major long-term cause of declines of species such as the products from the wild. However, the consensus tiger, across East and South Asia, poaching of tigers and among conservationists is that trade in farmed tigers other felids is the important short-term threat to sur- is likely to stimulate demand and provide the means vival of populations (Mills and Jackson 1994; Hemley to launder tiger parts derived from poaching, thereby and Mills 1999; Rabinowitz 1999; Wingard and Zahler reversing recent successes in reducing the magnitude 2006; Damania et al. 2008). Bones of other felids of the trade (Hemley and Mills 1999; Gratwicke era/. (clouded leopards, leopards, snow leopards, and lions) 2007; Nowell and Xu 2007). A recent survey are also used as substitutes for tiger bone and the trade measured attitude towards consumption of tigers in may therefore also impact populations of these species six Chinese urban areas. Of 1880 respondents, 43% (Wingard and Zahler 2006). The major Chinese invest- had consumed some product alleged to contain tiger ment and engagement in Africa may present a new products. The results indicated that while urban Chi- threat to lion populations, as it is nearly impossible to nese people are generally supportive of tiger conser- distinguish between lion and tiger bones. vation, there is a huge residual demand for tiger Trade in tiger parts, particularly bone, reached epic products which could potentially be filled by supply- proportions in the 1980s and 1990s, decimating wild ing parts from both wild and farmed tigers if the ban populations. For example, South Korean customs on trade in tiger parts is lifted in China (Gratwicke records from before South Korea acceded to CITES era/. 2008b). show that 8951 kg of tiger bone were imported from 1970 to 1993, with around half the bone derived from Indonesia. There has since been a concerted effort to control trade in tiger bone. Efforts have Management of trade and exploitation focused on international legislation, raising public The Convention on Biological Diversity (CBD) explic- awareness, dialogue with TAM specialists and users, itly recognizes sustainable utilization as a component and a commitment among TAM specialists to search of conservation of biodiversity and ecosystems (Con- for alternative medicinal products to replace the use vention on Biological Diversity 2003). The key to 194 Biology and Conservation of Wild Felids sustainable utilization is that use is responsibly man- other felid populations, such as trade in Asian cat aged and that populations are well monitored to en- skins. If the downturn in demand seen for spotted cat sure that overexploitation does not occur. However, skins is to be experienced for the Asian cats, a combi- sustainable utilization is often challenging in situa- nation of international and national legislations, con- tions where there is an inequity in distribution of sumer awareness, and education and engagement wealth, weak environmental controls, apathetic na- with the TAM industry is a first step in seeking to tional and international legislation, and endemic cor- reduce trade in tiger and other felid bones (Hemley ruption. Smith et al. (2003) found that poor and Mills 1999). The importance of both strictly governance, corruption, institutional failure, and so- applied legislation and public education is borne out cial and economic upheaval reduce the likelihood of by the decline in use of felid parts in China, where a successful conservation of biodiversity. Overexploita- concerted effort has been made to eliminate trade and tion of wildlife resources often goes hand in hand with raise awareness (Nowell and Xu 2007). uncontrolled habitat conversion (Rabinowitz 1999; Control of unsustainable trade in felids appears to Nyhus and Tilson 2004a) and poaching of tigers is depend on strong international controls, the willing- often linked to existing human-tiger conflicts, with ness and capacity of governments to implement and illegal trade being a by-product of conflict (Kumar and police conservation policy and provide disincentives Wright 1999; Johnson et al. 2006a). Household econ- to poaching and illegal trade, and the action of local omic and food insecurity has been shown to predis- and international conservation lobbyists and pres- pose people to illegal use of wildlife. In Zambia, sure groups. Consumer awareness and education improvement in food security reduced poaching and and engagement of stakeholders also appear crucial improved commitment to conservation initiatives in reducing or at least curtailing demand for felid (Lewis and Jackson 2005). products. Sustainable use of felid populations is clearly pos- sible, as evidenced by a sustainable fur trade in North America (Nowell and Jackson 1996). Key to this has been clear policies, effectively applied legislation, Conclusions and population monitoring and research. These con- ditions do not exist in the developing world. Trophy The world's ever-growing human population has hunting has the potential to be highly sustainable dramatically degraded most natural ecosystems, given its capacity for high revenue generation and causing an extinction event unprecedented in the protection of habitats and populations, provided in- last 65 million years. Large carnivores are the first stitutional mechanisms are in place to assure its sus- species to disappear under the onslaught of human tainability. It is also governed by international trade populations (Woodroffe 2001), having been eradi- agreements, such as CITES, and through government cated first from Europe, then decimated by Eur- conservation institutions. Also, sport hunters as a opeans in North America and more recently in Asia demographic grouping are often active in promoting and Africa. Today they persist only in large protected the conservation of the ecosystems they utilize. areas and in diminishing numbers in the least Trade in cat furs and the medicinal trade are less easy human-dominated of natural ecosystems. Barring a to control through international trade restrictions. sudden and unlikely reversal in human population However, systematic commercial hunting for spotted trend, the march towards expanding economies, or cat skins has seen a steady decline since the imposition attitudes towards the natural world, persistence of of trade restrictions by the United States, EU, and the larger felid species into the next century is a CITES, indicating that international action can have challenge we must all face. The outlook is dire for a positive impact. In addition, consumer awareness those who cherish wild cats. Smaller felids that do and sensibilities surrounding use of cat skins greatly not conflict with man and are not commercially reduced demand and aided the demise of spotted cat overexploited are likely to persist only where their trade in the Southern Hemisphere. This example may habitats are not deforested, overgrazed, or converted provide a template for controlling overexploitation of to agriculture. People and wild felids: conservation of cats and management of conflicts 195

The threats posed by humans to large felids are slow process compared to the short time left for some soluble if people sharing their landscapes have either wild tiger populations. economic or ethical incentives to preserve them. Like In contrast, African lions are declining largely due the ban on ivory trading that saved African elephant to continuing poverty and cultural modernization in populations to date, the successful ban on trade in Africa. At a time when increasing numbers of tropical spotted cat skins shows that conservationists impoverished people and their livestock are devastat- can have a significant influence on international mea- ing semiarid ecosystems, pastoralist people are also sures to preserve economically important species in abandoning traditional and effective methods of the developing world. Moreover, demand was reduced livestock protection in favour of agricultural pesti- when wearing spotted cat furs became socially unac- cides that allow them to eliminate entire populations ceptable, suggesting that moral pressures on society of predators at very low cost. Again, a major factor is may be equally effective. However, the continuing ineffective law enforcement due to lack of resources, decline of two of the most iconic felid species, the lack of interest, and pervasive corruption. However, tiger (Chapron era/. 2008a; Damania era/. 2008) and while tigers are poached because their parts are so lion (IUCN 2006b), provide a lens through which to valuable, lions are speared, trapped, and poisoned view the challenges facing conservationists when un- largely because they lack any economic value to the derlying conservation threats are not readily subject to humans that share their habitat. While both tourism legal or social measures. and sport hunting are highly lucrative and that Although conflict with people, habitat loss, and money could potentially be used to offset the costs reduced availability of wild prey are a significant of livestock depredation, very little profit from tour- problem for some tiger populations, poaching to sup- ism and hunting reaches the people whose precious ply markets for traditional medicine and skins in Asia livestock are killed by lions. Thus lions and other poses the greatest threat to the species (Dinerstein wild animals are nothing but an expensive nuisance et al. 2007; Chapron ef al. 2008a). It is unlikely that to rural people, with the inevitable result that they the trend will reverse in the absence of effective legal are being eradicated. Unless pastoralists and other measures to control trade and the sort of moral pres- rural Africans can earn significant income from wild- sure that made wearing of spotted cat furs unpopular life, there is no reason for them to conserve it. Inter- in the West. Intransigent and the dic- national fascination with these charismatic animals tates of fashion, increasing prosperity, and cultural gives them potentially great economic value. How- inertia in Asia, ensure an increasing and unprece- ever, this value is largely inaccessible to the rural dented demand by consumers for skins and bones. people who bear the brunt of living alongside large Meanwhile, pervasive corruption at all levels defeats carnivores. Big cats will survive in the wild only if legal sanctions. Demand for tiger products can only they become more valuable alive than dead to the be reduced through education about alternatives and people who share their landscapes. development of a sense of personal responsibility Conflict between people and large carnivores has for nature and wildlife and improved practice in wild- been a consistent theme throughout human history. life conservation. Unless consumers can be convin- As human populations burgeon, increasing pressure on ced that wildlife is more valuable roaming in forests remaining populations of carnivores will occur. Our than dismantled in apothecaries and upscale homes, understanding of the processes and patterns of use tigers and many other species seem doomed. How- and conflict may determine our ability to mitigate over- ever, it seems unlikely that such a shift in attitude can exploitation and conflicts and ensure survival of preda- be imposed by Western conservationists. It will only tory carnivores in a world increasingly dominated by accompany change in deeply held cultural values, a the needs and aspirations of the human species.