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
Wild felids and people have a complex and often paradoxical relationship. On the one hand, human- kind admires and reveres felids; cats appear as cultural icons and symbols across the ages. In addition, there is 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- trast strongly with the relationship between wild felids and people in areas where they coexist. Human con- flicts with wild cats, overexploitation of felid and prey populations, and habitat loss and fragmentation have
extirpated felid populations and still threaten many more. The ways in which people value and interact with organisms and their habitats is at the heart of conservation. This chapter explores some of the inter- relationships between people and wild felids, where human actions threaten felid populations.
Why conserve wild felids?
We preserve carnivores for aesthetic, symbolic, spiri- tual, ethical, utilitarian, and ecological reasons. Felids are culturally valued and are important as cultural icons and symbols. Felids are widely depicted in art,
162 Biology and Conservation of Wild Felids from Stone Age petroglyphs and cave paintings to more modern depictions of cats as art, reminding us that humans have interacted with felids for as long as we have been humans. Human culture is enriched by the symbolic use of felid images. Felids adorn many currencies, are heraldic symbols on coats of arms and appear widely on the badges of national sports teams.
They were revered by many cultures, for instance, the lion-headed goddess Sekhmet represented war and disease in ancient Egypt. The jaguar (Panthera onca) features significantly in Central American cultures.
Similarly, the tiger (P. tigris) is culturally important in South and East Asian cultures (Weber and Rabino- witz 1996).
Perhaps because of their place in our consciousness and the ease with which they are recognized, felids are used as umbrella and flagship species to conserve hab- itat, benefiting both felids and biodiversity generally.
Large carnivores are often highly mobile and viable populations require large tracts of suitable habitat and adequate prey populations. Thus, strategies for the protection of large felids also offer protection for 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 is seldom seen and outwardly unremarkable. Exam- ples of this include the protection, through an initia- tive called Paseo Panthera (the Path of the Panther), of biological corridors on the Panamanian Isthmus, used by jaguars, pumas (Puma concolor), and other wildlife.
'Tiger Reserves' protect significant areas of biodiversi- ty in India, as well as halting the decline of tigers, and tigers are used as a flagship to promote conservation efforts along border regions in the Indian subconti- nent and Indochina (Weber and Rabinowitz 1996;
Rabinowitz 1999; Seidensticker et al., Chapter 12, this volume; Sunquist and Sunquist 2002).
Moral philosophers and animal welfarists are united in their arguments that every species has an intrinsic conservation value that implies it has a right to survive. Intrinsic conservation values have served as an impetus for conservation. However, in- trinsic value in itself has been insufficient to secure successful conservation because humans are moti- vated more by economic self-interest than by ideas (Kellert et al. 1996). Economic benefits can provide powerful incentives to local communities to protect
biodiversity. Felids are economic assets and, when used sustainably through tourism, trophy hunting, or commercial exploitation can contribute substan- tially to both their own conservation and that of their habitats. For example, African lions (Panthera led) earned Amboseli National Park, Kenya, US
$27,000 per lion per year in tourism revenues (West- ern and Henry 1979). Trapping of furbearers or un- regulated trophy hunting can lead to controversial declines in populations. However, where it is carried out on a well-managed and sustainable basis, it can provide incentives for long-term protection, a good example being sustainable harvests of Canada lynx (Lynx canadensis) furs. Finally, felids play an impor- tant and often regulatory role within the ecosystems they inhabit (Mills and Biggs 1993; Karanth and Stith 1999; Grange and Duncan 2006). This ecologi- cal role needs to be factored into any valuation we make of felids in the wild. It must also be included in any assessment we make in valuing our relationship with nature and the pristine habitats that should form the baseline for all conservation efforts.
How people impact felids: anthropogenic threats to felid populations
Habitat loss is a global phenomenon, affecting all species. The earth's human population has increased from 3 billion to 6 billion since 1960. The global economy has increased sixfold and food production increased by 2.5 times. Nearly 25% of the earth's terrestrial surface is now under cultivation. The past 40-50 years have seen a sharp increase in the amount of land converted to agriculture, and projec- tions suggest that further conversion is to be expected in the future (Millennium Ecosystem As- sessment 2005). Conversion of natural habitat to agricultural land, urban development, and destruc- tion or fragmentation of habitats through logging, building infrastructure (e.g. dams, roads, power lines, oil, and mineral extraction), and other human activ- ity has serious impacts on wild felid populations.
Equally important is loss of prey populations through over-hunting, retaliatory persecution, habi- tat loss, or fragmentation. Predator population den- 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
overexploitation, such as the bushmeat trade, or habitat destruction) causes linked declines in felid populations (Karanth etal. 2004c; Henschel 2007).
Against a backdrop of global habitat loss, felids face a number of proximate anthropogenic threats.
One of the most important ways in which people impact felids is through increasing rates of non-nat- ural mortality (Fig. 6.1). For felid populations below their carrying capacity, anthropogenic mortality is thought to be largely additive rather than compen- satory to natural levels of mortality (Lindzey et al.
1992). By contrast, in highly protected populations anthropogenic mortality is rare, for instance only one (7%) of 14 leopard (Panthera pardus) deaths re- corded in Kruger National Park, South Africa, by Bailey (1993) was due to poaching; the rest were due to intraspecific fights, predation, and starvation.
In general, anthropogenic mortality in many studied felid populations is high. While this may in part be due to the fact that conservation biologists choose to study populations that are under threat, it is also an indicator of the impact that humans have on felid populations. Common sources of anthropogenic mortality are legal hunting and trapping, poaching, problem animal control (both legal and illegal), and vehicle accidents. Furthermore, felid populations
can also be exposed to diseases carried by domestic carnivores, often as a result of human encroachment into wild habitats. In some populations, anthropo- genic mortality can be extremely high and lead to population declines. For instance in Laikipia, Kenya, 17 of 18 tagged lions which died were killed in retri- bution for livestock raiding, with the population declining by about 4% per annum (Woodroffe and Frank 2005). Similarly, lion density was significantly reduced by conflict with local people on group ranches surrounding Masai Mara Reserve, Kenya (Ogutu et al. 2005). In a study of Amur Tigers, all seven recorded deaths of individually recognized adult females were due to poaching and 57% of cub mortality in this population was anthropogenic (Kerleyefa/. 2002, 2003).
Utilization of felid populations varies substantially in intensity, with mortalities from this source ranging from around 7% in Eurasian lynx (Lynx lynx) in Swit- zerland (Schmidt-Posthaus etal. 2002a) to 27 (62.7%) of 43 deaths of radio-collared lions around Hwange National Park, Zimbabwe (Loveridge etal., Chapter 11, this volume). Trophy hunting made up between 11.7% and 50% of recorded mortality in pumas (Logan etal. 1984; Lindzey etal. 1988; Cunningham etal. 2001), 37.5% in radio-tagged leopards in Natal, 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 bobcats (L. rufus) in Mississippi and Maine, United 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 populations few demographic data are available and this is particularly true of some populations hunted and trapped for the fur trade (e.g. spotted cats in South America in the 1960s and 1970s).
Conflict with people can lead to high levels of mortality, particularly where predator eradication occurs. For instance, an average of 29 (range 10-42) lions are destroyed each year in farmland around Etosha National Park, Nambia, where they pose a threat to livestock, with 563 lions killed over a 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.
1988; Cunningham et al. 2001). Around 3500 pumas per year are killed by people (through sport- hunting, protection of livestock, and vehicle acci- dents) in the western United States (Papouchis 2004).
Poaching can contribute substantially to mortality within a population. Ferreras et al. (1992) found that 42% of deaths of Iberian lynx (Lynx pardinus) in Do- nana National Park, Spain, were due to illegal poach- ing. Poaching accounts for a significant proportion of tiger mortality in India and Russia (Kumar and Wright 1999; Miquelle etal. 2005a) and Kenney etal. (1994, 1995) suggest that even moderate levels of poaching over relatively short time periods (~6 years) can lead to massive population declines of up to 95% in this spe- cies. In the Russian Far East, Chapron et al. (2008b) suggest that tiger populations cannot recover if annual mortality rates exceed 15%. Finally, road accidents can also be significant sources of mortality in some felid populations. Forty-five per cent of ocelot (Leopardus pardalis) mortality in Texas (Haines et al. 2005) and 48% of recorded puma deaths in Florida were due to road accidents (Taylor etal. 2002).
In this chapter we identify two key areas where felid populations face anthropogenic threats. Firstly, we explore the way felids impact people's lives and liveli- hoods and in turn the way in which people retaliate.
We then consider the impacts of trade and overexploi- tation on the conservation status of felid populations.
In each section, we offer a synthesis of the potential conservation and management solutions.
Conflicts between felids and people
Key areas of conflict between people and felids are through depredation on domesticated animals or game species and, less frequently, when large felids kill or injure people. Livestock raiding or human deaths often lead to retaliatory killing of the felids responsible. We will discuss, in the sections below, the issues of livestock depredation and human kill- ing, the consequences for felid populations, and po- tential management solutions in situations where such depredations occur.
Consequences of conflict for felid populations
Loss of human life or livelihood provides the impe- tus for people to attempt localized or sometimes wide-scale eradication of predators. Historically, eradication of carnivores has been a state-supported priority, incentivized by rewards and bounties. From 1860 to 1875, 4708 tigers and leopards were reported killed in India (Boomgaard 2001). In the United States, over 1160 pumas were culled from 1987 to 1990 in an attempt to limit livestock losses in ranch- ing areas (Johnson et al. 2001b). Wide-scale lethal control of predators still occurs in some frontier areas. For example, Michalski et al. (2006a) found that 110-150 jaguars and pumas were killed over a 12-month period by professional predator hunters in southern Amazonia, Brazil.
Contemporary government policies regarding pred- ator management are generally more enlightened.
However, in areas where depredations are perceived to impact livelihoods, both legal and illegal retaliatory killing of felids can extirpate populations. Frank et al.
(2006) describe the decimation of the lion population over large areas of Kenyan Masailand, where lion pop- ulations had previously been secure. The increasingly negative attitudes to wild predators in Masailand are at least nominally due to livestock depredation. How- 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,
coupled with ready access to agricultural poisons, may also be motivations behind increasing levels of illegal predator control. In Kenya, today, the use of agricul- tural insecticide Furadan (carbofuran) to kill predators is probably the single greatest source of lion mortality;
in Laikipia and Kajiado districts alone, a minimum of 70 lions are known to have been poisoned since 2001 (L.G. Frank, S. Maclennan, and A. Cotterill, unpub- lished data) and populations of vultures and other scavenging birds are plummeting throughout the country (S. Thomsett, personal communication).
Predation by felids on ungulate game species
Large felids can come into conflict with people over competition for wild prey species. Predators were even 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).
A worldwide meta-analysis of human-predator con- flict found that predators are reported to kill from 0.02% to 2.6% of livestock, but 9% of game species annually (Graham et al. 2005). Where wild ungulates are utilized by people for either commercial gain (com- mercial hunting and game farming) or for leisure (sport hunting), competition and conflict may occur between users and large felids. For example, cheetahs in Namibia kill economically valuable wild ungulates on commercial game farms, leading to persecution of the cats by farm owners (Marker era/., Chapter 15, this volume). Similarly, hunters in the Russian Far East come into conflict with tigers over real or perceived competition for ungulates (Miquelle et al. 1999a).
European lynx are persecuted when they compete with sport hunters for prized ungulate trophy species (Breitenmoser etal., Chapter 23, this volume).
Livestock depredation by felids
Felids are obligate carnivores. Humans and felids come into conflict in ecosystems, where a high proportion of ungulate biomass is made up of domesticated species.
For the most part, the smaller felid species do not cause
economically significant losses to domestic livestock, although species such as serval (Leptailurus served), wild- cat (Fells silvestris), and guina (Leopardus guigna) may occasionally prey on small stock or poultry (Sanderson et al. 2002c; Sunquist and Sunquist 2002). Generally, species ranging in size from the caracal (Caracal caracal;
~12 kg) to the size of the tiger (~235 kg) have been found to be the most problematic stock raiders. With the exception of the caracal and Eurasian lynx (~23 kg) all of the most import livestock-killing felids are 50 kg or heavier (Inskip and Zimmermann 2009). Table 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- ies, and reporting of widely different parameters and data. This makes rigorous evaluation of trends diffi- cult; however, there are a number of broad patterns that emerge and these are discussed in more detail in the following text.
Patterns of livestock depredation
Different size classes of livestock are vulnerable to different suites of felid predators. Smaller felids, such as lynx and caracals, prey on the smaller size classes of livestock such as sheep and goats, while the largest felids prey on the full size range of livestock. Interme- diate-sized felids, such as pumas and leopards prey on smaller livestock and juveniles of the larger species.
The spatial distribution of livestock depredation is often uneven. Stahl etal. (2001a, b) found that in the French Jura Mountains, there were consistent 'hot spots' of lynx predation on sheep. The areas were often also areas with high roe deer (Capreolus capreo- lus) densities, suggesting that lynx populations may have been sustained by reservoirs of natural prey.
Similarly, in southern and central Brazil, livestock were most vulnerable to puma and jaguar predation 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- stock predation by lions and leopards is often higher close to the boundaries of protected areas (Fig. 6.2).
This was found to be the case adjacent to Khutse Game Reserve, Botswana (Schiess-Meier et al. 2007) and Chirisa Safari Area, Zimbabwe (Butler 2000). In
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170 Biology and Conservation of Wild Felids
Figure 6.2 A village headman displays the remains of a cow killed by lions, Tshlotsho Communal Land, adjacent to Hwange National Park. © AJ. Loveridge.
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 it is thought that increased human activity in sur- rounding fields may also inadvertently provide protec- tion for stock (Madhusudan 2003).
northern Cameroon, levels of lion depredation were inversely correlated with distance to Waza National Park (Van Bommel etal. 2007).
Temporal patterns of livestock depredation are often predictable and in many cases vary with levels of avail- able natural prey. In many areas of Africa, livestock depredation peaks during the season when natural prey is most difficult for predators to acquire. This is usually the wet or rainy season, when prey are in better condition and often more widely dispersed (Patterson et al. 2004; Bauer and Iongh 2005; Kolowski and Hole- camp 2006). Hemson (2003) found that in the Maka- gadigadi ecosystem, Botswana, livestock-raiding lions killed fewer livestock during periods when migratory wildebeest (Connochaetes taurinus) and zebra (Equus burchelli) were present in their ranges (Loveridge et al., Chapter 11, this volume). In Laikipia, Kenya, livestock
Economic impact of livestock depredation Impacts of felid depredation on livestock vary, de- pending on the scale of livestock ownership, husban- dry techniques, livestock type, stocking density, and density of predators. Small-scale, subsistence livestock owners are often disproportionately impacted by losses, in part because they may lack resources to pro- vide effective protection for their stock. Loss of even an individual animal to small-scale livestock owners has a proportionally higher impact on herds or flocks than the same loss to owners with more stock. For this reason, Ikeda (2004) found that small-scale yak (Bos grunniens) herders in Kanchenjunga Conservation Area, Nepal, were more heavily impacted by snow leopard (Panthera undo) predation on their herds than wealthier, medium- to large-scale owners.
