Lion Predation on Livestock and Native Wildlife in Waza National Park, Northern Cameroon

DOI 10.1515/mammalia-2012-0063 Mammalia 2013; aop

Pricelia N. Tumenta * , Hermen D. Visser , Jacco van Rijssel, Lana M ü ller , Hans H. de Iongh , Paul J. Funston and Helias A. Udo de Haes Lion predation on livestock and native wildlife in Waza National Park, northern Cameroon

Abstract: The feeding ecology of lions ( Panthera leo, associated with human population growth (Woodroffe Linnaeus 1758) was investigated in and around Waza and Ginsberg 1998 ). Recently this has become increas- National Park, northern Cameroon. Diet was determined ingly problematic and threatens the persistence of the using global positioning system (GPS) data of lion kill sites African lion (Panthera leo, Linnaeus 1758) in many regions (clusters of GPS location points) collected using radio col- (Tumenta et al. 2010 ) where they regularly succumb lars. Lions consumed 14 different prey species, with five to persecution in retribution for livestock depredation species that were either medium (50– 200 kg) or large (Woodroffe and Frank 2005 ). Among a list of require- ( > 200 kg) in size forming the bulk of kills. The western ments, lions require a large and stable prey base to meet kob (Kobus kob kob ) was the most common (23.5 % ) wild their energetic needs (Schaller 1972 ). Lion diet thus varies prey of lions, but worryingly livestock (predominantly cat- across a range of predator, prey and environment-related tle) constituted as much as 21.6 % of the diet. This creates variables (Funston et al. 2001 , Radloff and Du Toit 2004 ). a conservation problem resulting in lions being killed in Lions prey on a broad range of species, typically ranging retaliation. However, this conflict is strongly exacerbated from medium to large-sized mammals. Lions in East and by herders driving their cattle into the park for forage and Southern Africa select a higher proportion of large prey water. Wild prey was also consumed outside the park (Hayward and Kerley 2005 , Owen-Smith and Mills 2008) (6.7 % ), suggesting excursions beyond the park boundary. than lions in West and Central Africa (Breuer 2005 , Bauer Lions showed a preference for wild prey over livestock et al. 2008 ). Typically the preferred prey weight of lions when relative abundances were considered. Management ranges from 190 to 550 kg, although the median weight efforts to reduce livestock intrusion into the park and of 350 kg is more selected (Hayward and Kerley 2005 ). to reverse the declining trends of wild prey populations Prey species outside the preferred weight range are gene- would significantly reduce predation of livestock and pre- rally avoided. However, lions are generally thought to be vent persecution of lions in this park, but conflict mitiga- opportunistic in their feeding (Schaller 1972 , Hayward tion measures outside the park are also urgently needed. and Kerley 2005 ) and to consume prey according to availability (Mills and Shenk 1992 , Breuer 2005 , Bauer et al. Keywords: African lion; diet composition; GPS data clus- 2008 ). Prey preference is influenced by many factors, ters; prey preference. notably prey size, prey availability (density and scarcity) and prey vulnerability (Schaller 1972 , Carbone et al. 1999 , *Corresponding author: Pricelia N. Tumenta , Institute of Hayward and Kerley 2005 ). Environmental Sciences, Leiden University, PO Box 9518, Predators have been reported to increase their attacks 2300RA Leiden, the Netherlands; Department of Forestry, University on livestock as a consequence of low natural prey avail- of Dschang, PO Box 479, Dschang, Cameroon, ability (Mishra 1997 , De Iongh et al. 2004 , Woodroffe e-mail: [email protected] and Frank 2005 ). Hemson (2003) found that lions in the Hermen D. Visser, Jacco van Rijssel, Lana Mü ller, Hans H. de Iongh and Helias A. Udo de Haes: Institute of Environmental Sciences, Makgadikgadi National Park, Botswana, increased the Leiden University, PO Box 9518, 2300RA Leiden , the Netherlands frequency of livestock predation during seasonal periods Paul J. Funston: Department of Nature Conservation , Tshwane of low wild prey availability. Patterson et al. (2004) also University of Technology, Private Bag X680, Pretoria 0001, South Africa recorded that livestock depredation increased when native prey was less accessible in Tsavo National Park, Kenya. Prey vulnerability is another factor that influ- Introduction ences prey selection. Livestock lack the physiological or morphological adaptations to escape from predators and Efforts to conserve large terrestrial carnivores are chal- defend themselves. They are easier to catch than wild prey lenged by increasingly complex requirements broadly and are thus considered vulnerable.

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The wild prey base for lions in Waza National Park, Observing lions on kills in low prey density areas with northern Cameroon, has drastically declined in recent intense human pressure is difficult. Lion diet was thus decades in response to a severe rinderpest outbreak in investigated using the method of GPS cluster searches 1982/1983, impact of the Maga dam on the natural regime of for carcasses of prey consumed by lions (Anderson and the Waza Logone floodplain, drought in the 1980s (Scholte Lindzey 2003 , Sand et al. 2005 , Tambling et al. 2010 ). Five 2005 , Scholte et al. 2007 ) and recently intensive human lions from four different “ prides ” were radio-collared from encroachment and poaching (De Iongh and Bauer 2008 , 2007 to 2009 using Vectronic GPS PLUS collars equipped De Iongh et al. 2009 , Foguekem et al. 2010 , Tumenta et al. with a very high frequency transmitter (Vectronic Aero- 2010 ). The lion population in Waza National Park has also space GmbH, Berlin, Germany) and African Wildlife Track- suffered persecution from herders for livestock depreda- ing GPS GSM collars (African Wildlife Tracking, Pretoria, tion and coupled with other factors declined from about South Africa). GPS positions were regularly downloaded 50 individuals in 2002 to fewer than 20 in 2008 (Tumenta from the collars. The locations of clusters of lion positions et al. 2010 ). Aerial counts of mammals in 2007 revealed a were entered into a handheld GPS unit, and the sites were minimum of 3466 herbivore mammals in the park against a visited to investigate the presence of kill remains. minimum of 21,000 livestock counted in a 5-km peri pheral Lions are most active during the night and feed on zone east of the park (Foguekem et al. 2010 ). average 4– 5 h when a kill of reasonable proportion is Thus, given the low wild prey densities, we expected acquired (Schaller 1972 ). Therefore, we selected nocturnal that lions in Waza National Park might switch their diet to GPS position clusters of 3 h and longer that were within a the more abundant livestock, considering its profitability radius of 50 m. The GPS point in the middle of the cluster as a large, vulnerable prey. Furthermore, because of the was put in a GPS unit and the location was visited by car or low wild prey densities, we additionally expected that on foot. At the location of the selected fix, carcasses were lions would be less selective and would consume a wider searched for within a 100-m radius. The location, species range of prey species. To test this, we assumed that the and size (small, < 50 kg; medium, 50– 200 kg; large, > 200 proportions of prey species carcasses identified at global kg; Bauer et al. 2008 ) were noted. Mean adult female body positioning system (GPS) clusters would reflect the actual weight (Kingdon 1997 ) was used for biomass calculations proportions of wild and livestock prey in the park. of prey. The age of carcasses was roughly estimated in the field and was later compared with the exact locations of GPS-collared lions. Materials and methods Prey selection was calculated using frequency of occurrence expressed as percentages. To determine prey Waza National Park lies between latitudes 10° 50 ′ and consumption based on body size, prey type, location and 11° 40 ′ and longitudes 14° 20 ′ and 15° 00 ′ . The climate is season, χ 2 goodness of fit test was performed. Prey pref- Sudano-Sahelian and semi-arid tropical. Rainfall is low erence was calculated using the Jacobs’ index with log Q and irregular between years, with a mean annual rainfall of giving positive values for preference and negative values 600 mm (Beauvilain 1995 ). Temperatures range from 15° C for avoidance (Jacobs 1974 ), with the formula D = (r-p)/ (January mean minimum) to 48 °C (April mean maximum). (r+ p-2rp), where r is the proportion of GPS-pointed kills for Waza National Park is characterized by three seasons: a a particular species (the fraction of a species in the diet) wet season, from June to October; a cold dry season, from and p is the proportional availability of that species (the November to February; and a hot dry season, from March fraction of the species in the habitat). Prey preference was to May. The park has three main vegetation zones: (1) The calculated only for species for which abundance estimates floodplain zone in the eastern half is dominated by heavy were available. Data of prey abundance were derived from cracking clay soils (vertisols) and grasses like Sorghum large mammal counts conducted in Waza National Park in arundinaceum , Pennisetum ramosum , Echinochloa pyrami- 2007 and 2010 (Foguekem et al. 2010 , Saleh 2010 ). dalis , Oryza longistaminata , Hyparrhenia rufa and Vetiveria nigritana . (2) The Acacia zone lying between the floodplain and the woodland is also a zone with clay soils, dominated by Acacia seyal trees interspersed with Balanites aegypti- Results aca , Piliostigma reticulatum and S. arundinaceum (3). The woodland zone found west of the park is on sandy soils The GPS cluster data revealed a total of 14 prey species and is dominated by Sclerocarya birrea , Anogeissus leio- from 162 carcasses of prey species identified. Prey species carpus and Lannea humilis (Wit 1975 ). included the western kob (Kobus kob kob), topi (Damaliscus

Brought to you by | Universiteit Leiden / LUMC Authenticated | 132.229.210.26 Download Date | 3/6/13 1:28 PM P.N. Tumenta et al.: Prey selection by African lions 3 korrigum), roan antelope ( Hippotragus equinus), red- cold dry and wet seasons, but predation during these two fronted gazelle (Gazella rufifrons ), warthog (Phacochoe- seasons was significantly higher compared to predation rus africanus ), giraffe (Giraffa camelopardalis ), elephant during the hot dry season (χ 2 = 13.78, df = 2, p = 0.05). Live- ( Loxodonta africana ), marabou stork ( Leptoptilos cru- stock predation was significantly higher during the cold meniferus), shoats (sheep and goats), cattle (Bos primige- dry season than during the wet and the hot dry seasons nius indicus), horse ( Equus ferus caballus), jackal (Canis ( χ 2 = 17.2, df = 2, p < 0.005). The gazelle, warthog and all aureus ), reedbuck (Redunca redunca) and ostrich (Struthio species of livestock were mostly consumed during the camelus). Percentage contribution of each prey species to cold dry season, whereas the topi, roan and giraffe were the total diet of lions was determined. Five species made mostly consumed during the wet season. During the hot up the bulk (84 % ) of the lion ’ s diet, with the western kob dry season, mostly the western kob was consumed. (23.5 % ) being the species most often selected, followed A majority of the prey species (67.3% ) were consumed by cattle (17.3 % ), topi (16.7 % ), roan antelope (13 % ) and at night (χ 2 = 19.36, df = 1, p < 0.001). Prey species con- giraffe (13% ). In terms of numbers of carcasses located, sumed inside the park (81.5 % ) was significantly higher wild prey constituted 78.4 % of the lion ’ s diet, against than prey species consumed outside the park (18.5 % ) 21.6% contributed by livestock (χ 2 = 52.25, df = 1, p < 0.001). ( χ 2 = 64.22, df = 1, p < 0.001). Cattle constituted 18 % of prey The fraction of livestock (21.6% ) predated by lions repre- consumed inside the park. Significantly more wild prey sents 4 % of livestock herds around Waza National Park. was consumed inside than outside the park (χ 2 = 111.8, The most preferred prey species was red-fronted df = 1, p < 0.001). However, about 6.7% of wild prey preda- gazelle followed by warthog, whereas the most avoided tion took place outside the park. Inside the park, male species was roan antelope (Table 1 ). Most prey species and female lions consumed both wild prey and livestock, (48.8 % of total) consumed belonged to the large-sized whereas outside the park only the males consumed both prey category, followed by medium-sized prey (40.7% ) types of prey, and females consumed strictly livestock. and small-sized prey (10.5% ). Prey consumption was significantly different with respect to prey size (χ 2 = 39.59, df = 2, p < 0.001). Discussion and conclusion The highest number of prey species were consumed in the Acacia vegetation zone (38.3% ) followed by the wood- The GPS cluster technique of identifying carcasses proved land vegetation zone (34.6% ) and the floodplain vegeta- to be useful in determining lion diet in Waza National tion zone (27.2 % ). However, the proportion of prey con- Park. This method is commendable for research in other sumed within various vegetation zones of the park was areas with lions in order to provide data for effective con- not different significantly (χ 2 = 3.11, df = 2, p = 0.21). servation of lions. However, it is important to determine In all three seasons (cold dry, hot dry and wet), sig- whether the age of the carcass corresponds with the age nificantly more wild prey was consumed than livestock of the GPS cluster. For this reason, it is recommended ( χ 2 = 52.25, df = 1, p < 0.001). Irrespective of prey type (wild that only recent GPS cluster data should be used as this prey and livestock), the highest predation was recorded will provide the most accurate and reliable results. Fur- during the wet season followed by the cold dry season, thermore, the decomposition rates of carcasses may differ whereas the lowest predation was recorded during the under different environmental conditions. Thus, it is pref- hot dry season. Predation was not different between the erable to study and determine the decomposition rate of

Table 1 Prey preference of lions revealed by carcasses (n = 162) located from GPS clusters in Waza National Park during 2008 and 2009.

Prey Number of Abundance Body mass ¾ Body % of prey % Prey Jacobs ’ species carcasses in Waza mass species available index

Kob 38 1,562 69 52 25.68 6.45 -0.06428 Topi 27 848 113 85 18.24 3.5 0.139135 Roan 21 148 252 189 14.19 0.61 -5.40648 Gazelle 6 28 20 15 4.05 0.12 1.228893 Warthog 4 21 60 45 2.7 0.09 1.132813 Giraffe 21 604 815 611 14.19 2.49 -0.21672 Cattle 28 17,459 330 248 18.92 72.12 0.020167 Shoats 3 3,543 35 26 2.03 14.63 0.294821

Brought to you by | Universiteit Leiden / LUMC Authenticated | 132.229.210.26 Download Date | 3/6/13 1:28 PM 4 P.N. Tumenta et al.: Prey selection by African lions carcasses prior to the use of this method. Kill sites located area to the floodplain zone that is more accessible to poach- using GPS clusters are biased toward large prey; therefore, ers. These results highlight the urgent need for management in areas that lack large prey this method should be com- to rehabilitate the watering points in this zone to prevent bined with scat analysis. animals from moving out. Livestock predation was highest Prey proportions in the lion ’ s diet reflected more or less during the cold dry season, coinciding with the period the proportion of occurrence of wild prey species in Waza when migratory livestock are moved en masse into the sur- National Park, with a preference for medium to large-size rounding area for dry season grazing. Management should ungulates. Prey preference indices, however, were not dic- intensify surveillance of the park and its boundaries espe- tated by the relative abundance, with the most preferred cially during this period, as this can reduce livestock intru- species, namely, red-fronted gazelle and warthog, being sion into the park and curb livestock depredation (Bauer relatively less abundant than more common medium and et al. 2010 , Tumenta et al. 2013). The fact that 18 % of the prey large-sized species. The preference for warthog could be killed within the park was cattle is evidence of how prob- the result of the slow evasion rate shown by this species lematic intrusion into the park could be. The highest pre- and the fact that the lion hunts them inside their burrows dation took place at night, concurring with earlier studies (Druce et al. 2004 ). It is unclear why the red-fronted on livestock depredation in Waza National Park (Bauer and gazelle was preferred most. Kari 2001 , Van Bommel et al. 2007, Tumenta et al. 2013) and Of concern, however, was that livestock comprised elsewhere (Hemson 2003 , Ogada et al. 2003 , Patterson et al. as much as 21.6% of the lion’ s diet within and outside the 2004 , Frank et al. 2005 ). Male and female lions both moved park, confirming earlier studies on livestock depredation out of the park and both killed livestock while outside the in the Waza National Park area based on interview surveys park. Livestock, especially cattle, has become an important (Bauer and Kari 2001 , Van Bommel et al. 2007 , Bauer et prey for lions in Waza National Park. Fortunately though, al. 2008 , 2010 , Tumenta et al. 2013). This contribution by it is not preferred relative to abundance. This is probably livestock to the lion ’ s diet represents 4 % of livestock herds because of long-term persecution by humans and reflects around Waza National Park; higher than 2.4% reported an adaptive response by lions. The future of lions in this by Patterson et al. (2004) in ranches neighboring Tsavo park depends on the efforts by management to reduce live- National Park in Kenya. These findings are, however, stock intrusion into the park and to increase populations of comparable to 5 % losses reported in Zimbabwe ’ s Gokwe wild prey within the park. Thus, are the key ecological con- communal lands (Butler 2000 ). Cattle comprised 80% of siderations to reduce lion predation on livestock. There are the livestock consumed by lions around Waza National also many key livestock husbandry approaches and human Park, highlighting the lion’ s preference for large prey attitude shifts that should be further encouraged to reduce when hunting livestock (Kruuk 1980 , Mills 1992 , Karani conflict. Although only a small amount (6.7 % ) of wild prey 1994 ). However, with respect to the preference index, lions predation was recorded out of the park, it would be neces- showed the lowest preference for cattle, indicating that sary for future research to determine why wild prey move although they are abundant, lions prefer to hunt wild prey out of the park. species. This can most probably be attributed to the persecution that generally follows the consumption of livestock Acknowledgements: We are extremely grateful to our (De Iongh et al. 2009 , Tumenta et al. 2010 ). driver, Hamidou, and the park guides who helped with The Acacia vegetation zone of the park was the richest GPS position clusters search for carcasses. Financial sup- in prey species occurrence, and lions thus killed the highest port for this study was provided by the Institute of Envi- amount of prey in this zone. During the dry season, all ronmental Sciences, Leiden University, the Netherlands. except one watering point in this zone dried up during the research period, leading to an outflux of wild prey from this Received May 4, 2012; accepted January 15, 2013

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