Factors Affecting Lion (Panthera Leo) Home Range, Movement and Diet in Waza National Park, Cameroon

Factors affecting lion (Panthera leo) home range, movement and diet in Waza National Park, Cameroon.

H.D. Visser1*, L. Müller1, P.N. Tumenta1,2, R. Buij1,2, H.H. de Iongh1

1Institute of Environmental Sciences, Leiden University, P.O.Box 9518, 2300 RA Leiden, The Netherlands 2 Centre for Environment and Development studies in Cameroon, Maroua/Department of Forestry, University of Dschang, PO Box 479, Dschang, Cameroon

Abstract

Key words: lions, habitats, home range, diet, movement

INTRODUCTION pressure on the environment, lion habitat and prey decreased and lions became less abundant and locally All over the world large carnivore populations are threatened (Bauer & Van de Merwe 2004). Bauer and under pressure from the increasing human popula- Van der Merwe (2004) estimated 16,500 – 30,000 free tion (Fuller, 1995; Nowell & Jackson, 1996). Threats ranging lions in all of Africa. to large carnivores include habitat loss, declining Although lions are threatened all over Africa, the In- natural prey populations and commercial exploita- ternational Union for Conservation of Nature (IUCN) tion, but most large carnivores are killed by people has listed the lion as Vulnerable on the global Red that defend their livestock (Woodroffe, 2001; Bauer, List (Bauer et al., 2008), populations in Southern Nowell & Packer, 2008). This conflict with livestock and East Africa seem to be quite viable and lions ex- owners is caused by the fact that carnivores have to ist in high densities locally (Bauer & ven der Mer- kill other animals to feed. In areas where wild prey we, 2004; Nowell & Jackson, 1996). Eastern Africa species are being replaced by livestock the conflict is counts 11,000 lions and Southern Africa 10,000. In intense (Ramakrishnan, Coss & Pelkey, 1999, Saber- Central and West Africa only 1800 lions are left and wal et al., 1994; Seidensticker, Christie & Jackson., populations are fragmented and declining (Bauer & 1999). When carnivores attack livestock, measures Van der Merwe, 2004). The population of lions over from livestock owners to protect their property are the past two decades has reduced by approximately inevitable. Often these measures include retaliatory 30% (Bauer et al., 2008). Only a few lion populations killing of the carnivores (Seidensticker et al., 1999; remain in protected areas with the largest in Central Woodroffe, 2000). African Republic (300) and second largest in Cam- A carnivore species that has suffered from conflict eroon (260). with humans in particular is the lion (Panthera leo) In Cameroon lions exist in two areas in the north: The (Patterson et al., 2004). In history lions were abundant Bénoué Ecosystem and Waza National Park (Bauer from Eurasia to Africa (Turner & Antón, 1997). When & De Iongh, 2005). In Waza National Park (WNP), human population expanded and pressure on the en- the research area of this study, lion population has de- vironment increased, lions disappeared from all of creased to the brink of extinction (Tumenta et al., in Europe and remained only in sub-Saharan Africa and press; De Iongh et al., 2009). In 1962, the lion popu- a small part of Asia (Nowell & Jackson, 1996). Due lation was estimated a 100 (Flizot, 1962), in 2001 at to human population growth and resulting increasing 50 (Bauer et al., 2003) and in 2008 only 14-22 lions were left (Kok, et al., 2008). The decreasing numbers of lions in WNP are mainly attributed to the increas- *All correspondence to: H.D. Visser, Institute of Environmental Sciences, Leiden University, P.O.Box 9518, 2300 RA Leiden, ing human activity in and outside the park that results The Netherlands. in prey base depletion and lion-livestock conflict (De E-mail: [email protected] Iongh & Bauer, 2008). The lion-livestock conflict is 2 H.D. VISSER ET AL.

Table 1 Number of prey species at Waza National Park from 1962 – 2007. Table is modiﬁed from De Iongh et al. (2004) with addition of data from Omondi et al. (2007) Only the main prey species of lions are includedd in the table. O = numbers based on regular observations in the park; T = transect; W = waterhole count; A = aereal survey; - = number unknown; * = number heavily underestimated.

Jan- Dec- 1962 1977 1977 1988 1994 1995 1996 1997 1998 2000 2001 2002 2007 Species O T T O W W W W/T T W A Kobus kob 25000 21993 13238 1700 2062 2573 4559 4517* 7665 1246* 9548 14489 1562 Damaliscus kor- 20000 749 605 - 1682 842 2395 720* 2198 236* 4284 7068 848 rigum Hippotragus 4000 349 223 - 1166 126 498 295* 1331 572 3868 5055 604 equinus Gazelle - 147 10 >300 621 455 440 204* 525 56* 593 228 28 ruﬁfrons Phacochoerus 600- 675 200 - 38 3 61* 16* 298* 44* 575 47* 21 aethiopicus 700 Redunca arun- 2000 25 39 - 66 50 301 54* 70 9* - - dicum Kobus 200 - - - 0 0 0 0 0 0 - 0 defassa a well known problem all over Africa (Bauer et al., and activities also had an impact on the prey species 2008). Often lion habitats are surrounded by human of lions in WNP. All herbivore species have decreased settlements. Livestock forms a competition for wild drastically since 1965. In the 1960s WNP counted herbivores with reduction of the prey population as a 25,000 kobs, about 5000 in the 1990s and in 2007 result. Livestock is a relatively easy prey to catch and only 1500 in 2007. The same holds for topi, with with the intrusion of livestock into the living area of 20,000 in the 1960s, 2000 in the 1990s and less than lions, easy prey is presented. Lions kill livestock and 1000 in 2007. Less drastically is the decrease of the to protect their property, lions get killed by herdsmen. roan antelope population with 4000 in the 1960s, less Although livestock is relatively easy to catch and than 1500 in the 1990s and less than 1000 in 2007 present in high numbers in WNP, it is not preferred as (Table 1). The southern reedbuck and defassa water- food source (Van Rijssel et al., 2008). buck have become locally extinct since the 1960s (De The negative effects of increased human population Iongh, Bauer & Hamling, 2004). In the late 1970s a

30,000

Roan 25,000 Western kob Topi 20,000 r e b

m 15,000 u N

10,000

5,000

0 1960 1970 1980 1990 2000 2010 Figure 1 Counts of three common, resident antelope species in WazaY eNationalar Park, western kob (Kobus kob kob), topi (Damalis- cus korrigum), and roan (Hippotragus equinus) since 1962 to present. Data from Scholte et al. (2007), Bobo Kadiri et al. (1998) and Omondi et al. (2007).

30,000

Roan 25,000 Western kob Topi 20,000 r e b

m 15,000 u N

10,000

5,000

0 1960 1970 1980 1990 2000 2010 Year

Factors affecting Home range, Movement and Diet in lions 3 drastic decrease in herbivore numbers was caused by a body weight of about 181 kg to 227 kg for males the construction of the Maga dam in 1979 (Scholte, and 113 kg to 136 kg for females, this means that lions Saleh & Serge, 2007). This dam prevented inunda- feed normally on large prey (Schaller, 1972). The size tion from the Logone river east of the park with a of prey lions are capable of killing is dependent on the dramatic impact on perennial vegetation, livestock, group size of the lion. When hunting alone, lions kill fisheries and birdlife in the downstream area of this smaller prey and when hunting in a group they kill river (Bauer, 2003; Scholte et al., 2007; Loth, 2007). larger prey (Schaller, 1972). Reversibly group size is Other factors inducing the rapid reduction of the an- smaller in areas with lower prey densities (Bauer, De telope species were rinderpest transmitted from live- Iongh & Di Silvestre, 2003). Small prey is killed in- stock because of an increase in livestock-antelope cidentally or out of necessity and animals weighing contact. Poaching was not seen as factor contributing more than 1000 kg are said to be relatively safe from to the decrease (Scholte et al., 2007). In the 1990s lion predation (Schaller, 1972). In an analysis of 32 and the beginning of 2000s populations of kob and different studies about lion diet Hayward & Kerley topi recovered little possibly by the (re)flooding of (2005) showed that lions prefer prey species within the floodplain (Scholte, 2007). In the 2000s this re- a weight range of 190-550 kg. The most preferred covery came drastically to an end and populations of weight was 350 kg, with gemsbok, buffalo, wilde- kob, roan and topi decreased again, with historic low beest, giraffe and zebra as favorite prey species. numbers of kob (Figure 1). Reason for this decrease Lions prefer prey species in a certain weight is an explosive increase in illegal activities inside the class, but what they actually eat is another thing. Be- park due to a breakdown of management (Tumenta et side prey preference many factors influence what a al., in press). lion eats. Most important are the characteristics of To get an idea of the lion-livestock conflict in WNP, the prey population: prey size, availability and den- it is important to gather ecological data of the lions. sity and scavenging (Schaller, 1972). Despite prefer- In this study we focus on two important factors that ences species that are not available can’t simply be might influence the lion-livestock conflict: sex and hunted. Ratios of prey occurrence in the environment habitat type. For these two factors, three important are more or less reflected in the diet (Schaller, 1972). variables are compared: diet, movement and home In Central and Western Africa large prey are hardly range. Previous studies have shown that these factors available. Lions in this part of Africa feed on smaller follow strong fluctuations over the seasons in WNP prey than the preferred size (Bauer et al., 2008). 49% (Bauer & De Iongh, 2005; Van Rijssel et al., 2008). of prey of the lions from Central and Western Africa We also included these fluctuations in our study. is medium sized (e.g. <200 kg) and 51% large (e.g. >200 kg), while in Southern and Eastern Africa 35% Diet is medium sized and 65% is large sized (Bauer et al.., To obtain the average daily energy requirements, a 2008). male lion needs to eat 7 kg and a lioness 5 kg (Schaller, In the African grasslands, the dynamics of herbiv- 1972). This food is obtained mostly by killing their ore populations depend mainly on food availability, own prey, but also partly by scavenging the kills of which on its turn depends on rainfall (Coe, Cumming other lions or other predators and the consumption of & Phillipson, 1976). Rainfall fluctuates over seasons animals that have died from diseases and other caus- and between areas. With these differences in rainfall, es. Lions are known to be opportunistic feeders that prey availability and density fluctuate over time and eat whatever they can catch (Schaller, 1972; Hayward space. Due to seasonal migration, herbivores avail- & Kerley, 2005; Eloff, 1984). As a result the diet of ability might differ over the seasons, or species might lions can differ between populations. Schaller (1972) not occur in a certain area at all. Prey density depends described lions in Kruger National Park that fed on as well on environmental changes between the sea- 38 different prey species and lions in Manyara Park sons. In the dry season herbivores are dependent on which fed on only 7 prey species. waterholes and their density can be very high locally Lions might be opportunistic in their feeding behavior on places with water and grass. In the rainy season and feed on a variety on species, but certain prey spe- as grass and water is everywhere, herbivore densi- cies are preferred above others. Generally carnivores ty is much lower (Smit, Grant & Devereux, 2007). of more than 21.5 kg feed on prey that is greater than Lion diet switches with these seasonal fluctuations of 45% of their own mass (Carbone et al., 1999). Lions herbivore availability and density (Scheel & Packer, are morphologically specialized in killing prey of their 1995, Ogutu & Dublin, 2002). In times of hardship, own size or larger (Sunquist & Sunquist, 1997). With lions might even eat species that are less preferred like 4 H.D. VISSER ET AL. springhare, honey badger and secretary bird (Eloff, Movement 1984). Another category of prey that lions might kill in times of hard conditions is livestock (De Iongh & Movement in animals is the result of the relation be- Bauer, 2008). tween characteristics of an individual (e.g. internal Livestock occur often in large densities in or state, motion and navigation capacity) and a broad close to lion habitat and form an easy to catch food range of external factors (Nathan et al., 2008). Gen- source. Because of the risk of getting killed by herds- eral goals for movement are gaining energy, seeking men, livestock is possibly not preferred (Van Rijssel safety, learning and reproducing (Nathan et al., 2008). et al., 2008). Nonetheless livestock is in some parts of In mammals energy requirements explain a substantial the world an important part of the diet of lions (Bauer variation in home range size and movements (Carf- & De Iongh, 2005, Van Rijssel et al., 2008). Several agno, 2007). Because movement is strongly related studies show that livestock predation by lions nega- to obtaining food, the above mentioned optimal for- tively correlates with wild prey density (Vijayan & aging theory (McArthur and Pianka, 1966) is appli- Pati, 2002; Hemson, 2003; Kolowsky & Holekamp, cable to movement as well. To gain the most energy 2005). Others suggest that livestock depredation by per time unit, an animal should limit the movement lions is reversibly induced by adverse circumstances invested in foraging as much as possible. Increased an example of which was described in WNP. Male li- distance traveled each day to find food has negative ons left WNP to feed on livestock exclusively (Bauer fitness consequences, decreasing the amount of en- & De Iongh, 2005, Van Rijssel et al., 2008). Because ergy invested in maintenance, repair and reproduction prey numbers are very low and decreasing, in WNP (Pontzer & Kamilar, 2009). lions possibly survive on a buffer of livestock (De On average lions spend about two hours a day Iongh & Bauer, 2008). In this study we try to test this walking, 40 to 50 minutes are spent eating and 20 to hypothesis. 21 hours a day lions are inactive (Schaller, 1972). A way to assess diet is applying optimal foraging This movement is not equally divided over the days, theory (MacArthur & Pianka, 1966). This theory pre- some days are spent without eating, while on some dicts the most optimal strategy (i.e. to be a special- days lions are eating 4 to 5 hours (Schaller, 1972). Al- ist or a generalist) for a predator using energy spent though movement occurs all day, lion activity peaks in searching and handling (i.e. pursuing, subduing are between 17:00 h and 8:00 (Schaller, 1972, Van and consuming it) prey species. The energy spent in Rijssel, 2008). Within these nightly hours movement searching and handling should be as small as possible is mainly related to hunting (Eloff, 1984; Schaller, and the energy intake as big as possible. Lions live 1972). Movement related to hunting ends when the normally in areas where prey is always near and don’t hunt was successful and a prey is killed, or, after an have to search long to make a kill. Handling the prey, unsuccessful hunt, when the temperature rises in the on the contrary, can be very time and energy consum- morning. In that way movement during hunting is ing. Applying optimal foraging theory implies for dependent on hunting success. On its turn hunting large predators as lions that the most profitable strat- success is dependent on a wide scale of environmen- egy is to specialize on the animals with the lowest tal factors (Funston, Mills, & Biggs, 2001). Because handling time (Begon, Townsend & Harper, 2006). hunting success is dependent on environmental fac- Lions living in an area with high prey abundance, can tors, and movement depends on hunting success, it is specialize on the most profitably prey species, but li- logical to expect that movement will show spatial and ons living in areas with low prey abundance should temporal differences. In an area with a high density of become more opportunistic and are forced to hunt suitable prey and good hunting conditions, lions will less profitable prey species. Of course there is a limit move less, than in areas with a low density of suitable to the capability of adaptation of lions to the lack of prey and bad hunting conditions (Ogutu & Dublin, suitable prey. Large carnivores simply cannot survive 2002). Since distance traveled to find food has nega- on a diet of species that are smaller than a certain size. tive fitness consequences (Pontzer & Kamilar, 2008), When hunting on too small prey, energy intake will the physical state of a lion might be deduced from the be lower than energy spent in searching and handling daily travel distance. As stated above, under sub-opti- prey (Carbone et al. 1999). Opportunism (e.g. the di- mal condition with low density of suitable prey, lions versity of prey species in the diet) could be a incicator might change their diet to livestock, an easier to catch for the state of the environment. prey (Bauer & De Iongh, 2005). In the surroundings of WNP livestock is available in large numbers. During an aerial survey 20,000 individuals of livestock were Factors affecting Home range, Movement and Diet in lions 5 counted near the borders of WNP (Omondi, Mayien- had larger home ranges in the wet season than in the da & Tschamba, 2008). Because livestock is easier to hot dry season (Schoe et al., 2007; Van Rijssel et al. catch, lions move less when they hunt livestock than 2008). In WNP home range sizes might also be in- when hunting on wild prey (Hemson, 2003). fluenced by differences in habitat qualities between In WNP prey densities differ per habitat type and show the forest zone and the floodplain, with bigger home seasonal fluctuations (Scholte et al., 2007). These ranges in the forest and smaller in the floodplain. Be- spatial and temporal fluctuations in food availability cause of strong seasonal fluctuations in prey abun- might result in a difference in habitat suitability for dance in the floodplain, differences over the season the lions. Because prey densities are lower in the for- might be bigger. est zone, this habitat type might be less suitable. This Male lions have also been shown to have larger home difference in habitat type might have an impact on the ranges than females (Schoe et al., 2007; Van Rijssel daily travel distance of lions. et al. 2008). These bigger home ranges are partly induced by the fact that males are bigger than females Home range and have higher energetic needs and partly by the territorial behavior they display (Lehman et al., 2008). According to Burt (1943) a home range is: “…that Because females with cubs stay close to their cubs, area traversed by the individual in its normal activi- they have smaller home ranges than females without ties of food gathering, mating and caring for young.” cubs. Home range size is dependent on a broad scale of factors. Between species from the order of the carnivores Differences between sexes home range size increases with metabolic needs (Git- tleman & Harvey, 1982). Servals (Leptailures serval) Between lions of both sexes, an obvious sexual di- have an average home range of 1.5 km2, leopards morphism exists. Adult males are almost twice the (Panthera pardus) 23.6 km2 and lions 240.0 km2 (Git- weight of adult females (Smuts, Robinson, & Whyte, tleman & Harvey, 1982). Carnivore home ranges are 1980) and have visually impressive a conspicuous generally as large as is necessary but as small as pos- manes. It is broadly assumed that females do most sible to satisfy energetic needs (Gittleman & Harvey, of the hunting in the pride, while adult males tend to 1982; MacDonald, 1983). Upper limits are determined refrain from hunting and pursue prey less often than by energy expenditure during territorial defense (Ber- females (Funston et al., 2001). Males obtain more of- tram, 1973) while lower limits are governed by food ten by scavenging from kills of female lions or other availability (Van Orsdol, Hanby & Bygott, 1985). predators. Males do participate in the hunt of large Other important factors influencing home range size prey. Funston et al. (1998) found that territorial males are habitat qualities such as waterholes and rivers, or and especially non-territorial males also hunt success- access to refuges (Spong, 2002). Furthermore home fully for themselves. Socially, females are the stable range size is directly related to body weight. With in- content of a pride and often only a part of their life creasing body weight, metabolic needs increase and nomadic, while males are only a relative short part therefore home range size (Harestad & Bunell, 1979, of their life part of a pride and the rest of their live Gittleman & Harvey, 1982). Social factors as group nomadic (Schaller, 1972). The differences between size and competition also influence home range size males and females have consequences for diet, home (Van Orsdol et al., 1985). Between lions home ranges range and movement. can vary substantially in size. Van Rijssel et al. (2008) Males are known to be more successful in made an overview of data available on home range hunting larger prey and females in hunting medium sizes and found home ranges ranging from only 22 sized prey (Funston et al., 2001, Funston et al., 1998). km2 to 7337 km2. In WNP large prey are less abundant than medium Because of spatial and temporal differences in sized prey (Scholte et al., 2007; Omondi et al., 2008) habitat qualities (Coe et al., 1976), home range sizes and males might encounter more problems in get- vary between habitats and seasons. Lion home range ting prey than females. Additional males have higher size and population density are exponentially corre- metabolic needs, because they are larger than females lated with prey abundance during time of prey scarci- (Schaller, 1972). This results in the expectation that ty (Hemson, 2003). In the dry season, when prey den- males will have larger home ranges than females. Be- sities are locally very high, home ranges are smaller sides metabolic needs, territorial behavior of pride than in the rainy season, when prey densities are low. males leads to larger home ranges (Lehman et al., In both WNP as Bénoué NP, North Cameroon, lions 2008). The expectation that males have larger home 6 H.D. VISSER ET AL. ranges than females is conformed by former research stock conflict which on its turn is at the base of the in WNP (Bauer & De Iongh, 2005; Van Rijssel et al., near extinction of the lions in WNP (Tumenta et al. in 2008). These factors (i.e. low density of large prey, press; De Iongh et al. 2009). We expect that the sex behavior) will imply that males move more than fe- of lions and the difference in habitat will result in dif- males. In this study we assume that nightly movement ferences in ecological factors and contribution to the is an indicator for hunting success, so we expect males lion-livestock conflict. move more than females during nightly hours. Males are expected to suffer more than females from Besides bigger home ranges and travel dis- the degradation of WNP. Indications for this differ- tances, the differences between males and females ence will be a bigger home range size, more move- will have an effect on diet. Usually males are more ment and differences in diet. Males are expected to specialized in hunting bigger prey than females. In contribute more to the lion-livestock conflict than fe- WNP males will have more difficulties to meet their males. energetic needs and as stated above, in times of hard- The forest zone is expected to be a less suitable habi- ship lions might switch their diet to livestock (De tat for lions, while the floodplain shows stronger fluc- Iongh & Bauer, 2008). We expect males to feed more tuations between the seasons. Indicators for this dif- on livestock than females. This hypothesis has been ference will be larger home ranges for lions from the proved by the findings of former research on other forest zone and bigger fluctuations for lions from the male lions in WNP (Bauer & De Iongh, 2005). floodplain, more movement for lions from the forest Differences between habitat types zone and bigger fluctuations for lions from the flood- Different habitat types differ in characteristics that are plain and more opportunism for lions from the forest needed for lion survival. Lions living in habitats with zone. Lions from the forest zone are expected to eat lower prey density have to search longer and walk more livestock, while lions from the floodplain are longer distances per day to find their food. To opti- expected to show stronger fluctuations. mize their chances, they need to increase their home range. Because in the lion diet the ratios of prey spe- MATERIALS AND METHODS cies in the environment are to some extent reflected (despite preferences), diet will differ between habi- Research area tats. In WNP, roughly three habitat types are present: a woodland zone, an acacia zone (combined in the This study was conducted in WNP, in the Extreme forest zone) and a floodplain. The forest zone on the North Province of Cameroon. The park covers an area higher parts of the park and has a lower density of of approximately 1600 km2 and lies between 11º00’- antelope species as kob, topi and roan, but contains 11º50’N and 14º30’-14º75’E (Figure 2). WNP has an the majority of all giraffes (Scholte et al., 2007). On average altitude of 300m in the East and 320m in the the floodplain, a fluctuation in prey densities is caused West, rising to 500m on top of the rocky inselbergs by the annual floods. From June to the end of October around Waza village in the West. WNP climate is Sou- water is available and animals are not dependent on dano-Sahelian, semi-arid tropical with three seasons: waterholes. So the woodland might be a less suitable the wet season from Jun till October, cold dry season environment, with low fluctuations in food availabil- from November till February and the hot dry season ity, while the floodplain is more suitable, but shows from March till may. Mean annual rainfall is 650 mm more fluctuations in food availability. In this study with a strong fluctuation between the years. During we see movement, home range size and livestock in the rainy season, the eastern part of the park becomes the diet as indicators for environmental suitability to flooded by water from the Logone river, 15 km east lions. For the forest zone we expect these factors to of the park. The higher part (south-west) of the park be more stable over the year and for the floodplain we remains unflooded, but waterholes fill up with water. expect more fluctuations. Livestock predation of lions During the dry seasons the water evaporates and the from the woodland might occur all year, while lions river withdraws, leaving only a few permanent water- from the floodplain will cause more problems in the holes at the end of the dry season. wet season and cold dry season. Vegetation Hypotheses The south-west of the park contains sandy soils with Home range, movement and diet are important eco- woodland savannah. Dominant tree species are Scle- logical factors that are interwoven with the lion-live- rocarya birrea, Anogeissus leiocarpus, Lannea humi- Factors affecting Home range, Movement and Diet in lions 7

lis, Combretum sp., Feretia apodenthera and Acacia (Phacochoerus africanus, Pallas 1676), reedbuck dudgeoni. The clay soils in the center of the park are (Redunca redunca, Pallas 1767) and Grimm’s duiker characterized by thorny shrub savannah dominated (Sylvicarpa grimmia, Linnaeus 1758). The latter two by Acacia sp., Balanites aegyptiaca, Pilliostigma re- species have become extremely rare. Large carnivores ticulatum, Calotropis procera and Ziziphus sp. These in WNP are lion (Panthera leo, Linnaeus 1758) spot- two vegetation zones are sometimes combined in the ted hyena (Crocuta crocuta, Erxleben 1777), striped “forest zone” (Bauer, 2003). The fl oodplain in the hyena (Hyena hyena, Linnaeus 1758). Besides mam- north west of the park is dominated grasses as Sor- mals WNP contains an impressive variety of birds ghum arundinaceum, Pennisetum ramosum, Echino- with ostrich (Struthio camelus, Linnaeus 1785) and chloa pyramidalis, Oriza longistaminata, Hyparrhen- black crowned crane (Balearica pavonina, Linnaeus aia rufa and Vetiveria nigritana (Scholte et al., 2007; 1785). Leopard (Panthera pardus, Linneaus 1833), Bauer 2003; De Iongh et al., 2004). cheetah (Acinonyx jubatus, Schreber 1775), water- buck (Kobus ellipsiprymnus, Ogilby 1833), bushbuck Herbivore diversity (Tregelaphus scriptus, Pallas 1776), and red fl anked duiker (Cehpalophu rufi latus, Gray 1846) have be- Waza National park is an important area for at least 30 come extinct the last few decades (Scholte et al., mammals. Herbivore species present in WNP are: Gi- 1999; Bauer, 2003). raffe (Giraffa camelopardalis, Linnaeus 1758), Ele- phant (Loxodonta africana, Blumenbach), red fronted Research lions gazelle (Gazelle rufi frons, Gray 1846), western kob (Kobus kob kob, Erxleben 1777), topi (Damaliscus In this study data of fi ve collared lions are used (Tu- korrigum korrigum, Burchell 1823), roan antelope menta et al., 2007; Tumenta et al., 2008). An over- (Hippotragus equinus, Desmarest 1804), warthog view of the characteristics of these lion is given in Table 2.

14.50 15.00 11. Nigeria

Floodplain Acacia zone

Woodland zone Chad Road Cameroon Inselberg Village Waza National Park

Figure 2 Map of WNP. 8 H.D. VISSER ET AL.

Table 2 General data for the collared lions. L1 L2 L3 L4 L5 Name Adam Jean Pierre Elisabeth Fanne Rosy Sex Male Male Female Female Female Vegetation Forest Floodplain Floodplain Floodplain Forest zone Data start 05-10-2007 05-05/2007 05-05-2007 05-06-2007 05-18-2008 Data end 04-04-2008 11-18-2007 06-12-2008 09-25-2007 02-01-2009 Days 331 198 405 143 260 Death Apr 2008 Nov 2007 - - Feb 2009

Home ranges GPS-clusters Calculation and mapping of the fixes and the home Lions hunt between sunset and sunrise and when ranges was done with ArcView 3.2 GIS (ESRI, 1992) the hunt is successful and they have killed a prey of with the extensions spatial analyst and animal move- reasonable size they eat 4-5 hours (Schaller, 1972). ment. Lion inactivity is visible in the dataset as a cluster of The dataset of the first four lions was too big GPS fixes on one place. Fixes were recognized as one to execute home range calculations with ArcView 3.2 cluster when they were less than 50 meters from each GIS. To reduce the dataset only one fix in four hours other. Clusters of 3 hours or more between 18:00 and was used, leaving only 6 fixes per day in stead of 48 6:00 were selected for further investigation. Of the fixes per day. Obvious outliers (some fixes were on fixes within the cluster, the average latitude and lon- Antarctica or even not on earth) were removed by vis- gitude were calculated. This value was put in a GPS ible inspection in Microsoft Excel. Home ranges were device (Garmin eTrex H) and visited by car. At ar- assessed as Harmonic Mean. The Harmonic Mean was rival on the location of the cluster, we searched with- calculated using the Kernel-test (cell size = 0.0009), in a circle with a radius of 100 meters for carcasses. with the 95% Harmonic Mean as the boundary of the When we found a carcass more than 100 meters away home range, the 50% Harmonic Mean as the core area from the centre of the cluster, it was not recognized of the home range and 5% the heart of the core area as belonging to the collared lion. Of any found we (White & Garrot, 1990). Home range calculations noted location, species, life stage (juvenile, sub-adult, were executed for all of the data and for selections of adult), age (recent, old, very old), sex, grass and shrub data from the hot dry season, the cold season and the cover of the direct surroundings and vegetation type. wet season. When we found a carcass that was older than the age of the cluster, the carcass was not recognized as be- Daily travel distance longing to the collared lion. Of each carcass a photos was made with a Canon Powershot SX100 IS photo For distance calculations, the same dataset was used camera. Prey species were divided in two weight cat- as for the calculations for home range. The calcula- egories (Bauer et al., 2008). A prey was scored as me- tions were executed using Microsoft Excel and no dium when <200 kg and large when >200 kg. Of each data reduction was needed. found carcass prey type (i.e. wild prey or livestock) was assigned. Data analysis This method was applied to all lions, except L4. Her data were very old and no carcasses, or very old and For the analysis of the data SPSS 16.0 was used. Sig- scattered, were found during fieldwork. The cluster nificance of differences in diet between sexes and be- search turned out to be successful for data less than tween habitat types was calculated with a Chi-square one year old. test. Significance of differences in distance and home The fieldwork for L1 and L2 was done in February range size between sexes and habitat types was cal- till June 2008 (Kok et al., 2008; Van Rijssel et al., culated with one-way ANOVA. Prey preference was 2008) and for L3 and L5 from February till June 2009 calculated with the method of Jacobs (1974). (Table 2). Factors affecting Home range, Movement and Diet in lions 9 Table 3 Overview of characteristics of L1, L2, L3, L4 and L5; Groups: M = males, F = females, FP = floodplain, WL = woodland, T = total, all data for groups are averages; Daily travel distances in km; Home range sizes in km2, 95%, 50% and 5% harmonic mean; Time spend outside the park borders in % of all the fixes; Prey type in % of all prey, W = wild prey, L = livestock;.

L1 L2 L3 L4 L5 M F FP WL T Daily travel distance All data day 1.05 1.22 1.61 1.44 1.91 1.11 1.67 1.47 1.41 1.45 night 6.80 7.16 4.60 4.02 3.26 6.93 4.08 5.17 5.27 5.22 total 7.84 8.39 6.21 5.46 5.08 8.05 5.72 6.64 6.65 6.64 Hot dry day 1.51 0.68 2.08 2.06 2.01 1.32 2.07 1.89 1.58 1.79 night 6.36 5.01 5.81 4.53 4.25 6.05 5.52 5.55 6.05 5.72 total 7.87 5.68 7.89 6.59 6.12 7.37 7.58 7.44 7.61 7.50 Wet day 0.83 1.37 1.21 1.31 1.65 1.10 1.39 1.29 1.23 1.27 night 9.18 7.53 4.20 3.91 2.40 8.36 3.50 5.29 5.85 5.52 total 10.01 8.90 5.41 5.22 4.02 9.46 4.88 6.59 7.06 6.78 Cold dry day 0.98 0.78 1.56 2.35 0.94 1.94 1.44 1.64 1.56 night 3.24 7.13 3.39 4.54 3.87 3.96 4.00 3.88 3.92 total 4.21 7.91 4.96 6.66 4.82 5.80 5.44 5.42 5.43 Home range size All data 5% 1,7 2,6 1,9 2,3 2,6 2,1 2,3 2,1 2,3 2,2 50% 27,5 84,2 63,8 82,6 134,8 55,8 93,7 81,1 76,9 78,6 95% 795,4 817,9 861,4 327,4 731,8 806,6 640,2 763,6 668,9 706,8 Hot dry 5% 1,5 1,4 1,9 1,7 1,5 1,5 1,7 1,5 1,7 1,6 50% 21,4 34,3 39,4 36,7 28,6 27,8 34,9 25,0 36,8 32,1 95% 150,7 126,4 240,7 143,8 244,0 138,5 209,5 197,4 170,3 181,1 Wet 5% 1,7 2,5 2,1 2,0 2,3 2,1 2,1 2,0 2,2 2,1 50% 32,9 171,3 31,8 50,6 82,8 102,1 55,1 57,9 84,6 73,9 95% 816,0 840,3 340,9 310,2 404,7 828,2 351,9 610,4 497,1 542,4 Cold dry 5% 1,4 1,4 1,6 - 2,0 1,4 1,8 1,7 1,5 1,6 50% 24,9 24,4 52,0 - 80,8 24,7 66,4 52,9 38,2 45,5 95% 449,9 371,3 566,2 - 694,1 410,6 630,1 572,0 468,7 520,4 Time spend outside park borders Total 26 20 19 0 1 23 7 14 13 13 Hot dry 12 0 6 0 0 6 2 6 2 3 Wet 19 22 1 0 0 21 0 10 8 8 Cold dry 42 25 52 - 3 33 28 22 38 24 Prey type Total W 79 70 78 96 75 85 70 84 80 L 21 30 22 4 25 15 30 16 20 Hot dry W 75 100 87 85 87 90 75 86 L 25 0 13 15 13 10 25 14 Wet W 81 64 100 92 72 96 78 85 81 L 19 36 0 8 28 4 22 15 19 Cold dry W 77 47 100 77 72 47 89 74 L 23 53 0 23 28 53 11 26 Prey diversity Total prey 41 46 30 26 87 56 76 67 143 87 Different species 10 9 6 10 19 16 15 20 14 19 Prey diversity 0,24 0,20 0,20 0,38 0,22 0,29 0,20 0,30 0,10 0,22

RESULTS od in 2009 and no new data were obtained since June 2008. The collar of L4 became disfunctional in No- L1 was killed in April 2008, 11 months after he was vember 2007. We observed her and she looked quite collared in May 2007 and L2 was killed in November vital and her collar was still in place (Table 3). 2007, 6.5 months after collaring. L1 and L2 both were killed outside the park border most likely by pastoral- Daily travel distance ists who were defending their livestock. L5 died in February 2009, 9.5 months after she was collared. Her Daily travel distances were calculated for all of the remains were found inside the park. Despite intensive ﬁve lions as presented in Table 3. For these ﬁve lions telemetry L3 was not found during the research peri- the period of investigation was different (Table 2). 10 H.D. VISSER ET AL.

50000 Season 50000 Season Hot Dry Hot Dry 40000 Wet 40000 Wet Cold Dry Cold Dry 30000 30000

Distance 20000

10000 10000

0 0 Forest zone Woodland Male Female Habitat type Sex Figure 4 Daily travel distance for lions from the forest zone Figure 3 Mean daily travel distance for males and females and lions from the woodland.

Within the period of investigation L1 traveled 7.84 4.77) and for woodland lions from 0.086 km to 33.89 km on average per day, L2 8.39 km, L3 6.20 km, L4 km (µ = 6.65 km, σ = 5.55). The difference between 5.46 km and L5 5.08 km. The five lions traveled the lions from different habitat types is not significant (F most in the hot dry season (µ = 7.50, σ = 4.88), less < 0.001, p = 0.99). Lions from different habitats show in the wet season (µ = 6.78, σ = 5.44) and least in the same pattern in daily travel distances over the sea- the cold dry season (µ = 5.43, σ = 4.26) (Table 3). sons (Figure 4). Lions from both floodplain and the This difference was highly significant (F = 12.849, p forest zone travel most in the hot dry season (resp. µ < 0.001). = 7.44, σ = 4.22 and µ = 7.61, σ = 5.99), less in the Not surprisingly for all lions more traveling took wet season (resp. µ = 6.59, σ = 5.12 and µ = 7.06, σ = place during the nightly hours, 5.22 km on average, 5.88) and least in the cold dry season (resp. µ = 5.44, than during the day, 1.45 km on average. σ = 3.94 and µ = 5.42, σ = 4.46). Lions from the floodplain and forest zone show hard- Difference between the sexes ly any difference in travel distance during daylight Daily travel distance for males ranged from 0.31 (resp. µ = 1.47 km, σ = 1.75 and µ = 1.41 km, σ = km to 41.43 km (µ = 8.05 km, σ = 6.44) and for fe- 1.58) and nightly hours (resp. µ = 5.17 km, σ = 4.12 males from 0.07 km to 20.33 km (µ = 5.72 km, σ = and µ = 5.27 km, σ = 5.18). This observation weighs 3.75). Males moved significantly more than females for all seasons. (F = 68.90, p < 0.001). Over the seasons, males and females show a difference in daily travel distance. Home range Males traveled most in the wet season (µ = 9.46 km, σ = 6.63), less in the hot dry season (µ = 7.37 km, σ Home range size = 6.05) and least in the cold dry season (µ = 4.82 km, σ = 4.92 m). Females traveled most in the hot dry Harmonic means (Kernel 5%, 50% and 95%) were season (µ = 7.58 km, σ = 4.04), less in the cold dry calculated for five lions as presented in Table 3, Figure season (µ = 5.80 km, σ = 3.78) and least in the wet 5,Figure 6,Figure 7 and Figure 8. The average home season (µ = 4.88 km, σ = 3.29) (Figure 3). range size (Harmonic Mean 95%) for all data of the Males move more than females during nightly hours five lions was 706,8 km2. Strong fluctuations over the (resp. 6.93 km and 4.08 km). This difference is highly seasons were observed. Home ranges were biggest in significant (F = 134.048, p < 0.001). On the contrary the wet season, 542,4 km2 on average, smaller in the females move more in daylight than males (resp. 1.67 cold dry season, 520,4 km2, and the smallest in the Page 1 km and 1.11 km). This difference is highly signifi- hot dry season, 181 km2. Page 1 cant as well (F = 36.048, p < 0.001). Both observation weighs for all the seasons (Table 3). Differences between sexes In the hot dry season and the cold dry season home Differences between habitats ranges of females (respectively 209 km2 and 630 Daily travel distance for lions from the floodplain km2) are on average larger than those of males (re- ranged from 0.07 km to 41.43 km (µ = 6.64 km, σ = spectively 138 km2 and 410 km2). This difference Factors affecting Home range, Movement and Diet in lions 11

L1 kernel 95%, 50%, 5% L2 kernel 95%, 50%, 5%

L3 kernel 95%, 50%, 5% L4 kernel 95%, 50%, 5%

L5 kernel 95%, 50%, 5% Roads Waterholes Villages Woodland zone Acacia zone Waza National Park Floodplain } Cameroon Nigeria Chad Kernel 95% Kernel 50% Homerange Kernel 5% }

Figure 5 Homerange calculated with a Kernel test (H=0.02) for all available data; HM (95%, 50% and 5%). 12 H.D. VISSER ET AL.

��

Figure 7 Homerange calculated with a Kernel test (H = 0.02) for data from the hot dry season; HM (95%, 50%, 5%). No data from the cold dry season are available for lion 4. Factors affecting Home range, Movement and Diet in lions 13

L1 kernel 95%, 50%, 5% L2 kernel 95%, 50%, 5%

L3 kernel 95%, 50%, 5% L4 kernel 95%, 50%, 5%

L5 kernel 95%, 50%, 5% Roads Waterholes Villages Woodland zone Acacia zone Waza National Park Floodplain } Cameroon Nigeria Chad Kernel 95% Kernel 50% Kernel 5%

Figure 8 Homerange calculated with a Kernel test (H = 0.02) for data from the wet season; HM (95%, 50%, 5%). 14 H.D. VISSER ET AL.

L1 kernel 95%, 50%, 5% L2 kernel 95%, 50%, 5%

L3 kernel 95%, 50%, 5% L5 kernel 95%, 50%, 5%

Roads Woodland zone Cameroon Kernel 95% Waterholes Acacia zone Waza National Park Nigeria Kernel 50% Villages Floodplain } Chad Kernel 5% Figure 6 Homerange calculated with a Kernel test (H = 0.02) for data from the cold dry season; HM (95%, 50%, 5%). No data from the cold dry season are available for lion 4. is not significant (F = 0.544, p = 0.458). In the wet Differences between habitats season males (828 km2) have a much bigger home In the hot dry season, the wet season and the cold dry range than females (351 km2). This difference is sig- season home ranges of forest zone lions (respective- nificant (F = 156.067, p = 0.001). Between the sexes ly 197 km2, 610 km2 and 572 km2) are bigger than sizes of home ranges show a different pattern over home ranges of floodplain lions (respectively 170 the seasons. Home ranges of males are biggest in the km2, 497 km2 and 469 km2). For both forest zone wet season, smaller in the cold dry season, and the lions and floodplain lions home ranges are the big- smallest in the hot dry season, while the home ranges gest in the wet season, smaller in the cold dry season of females are largest in the cold dry season, smaller and the smallest in the hot dry season. When the to- in the wet season and the smallest in the hot dry sea- tal dataset is considered, forest zone lions (764 km2) son. When the total dataset is considered, males (806 have a bigger home ranges than floodplain lions (669 km2) have a bigger home range than females (640 km2). This difference was not significant (F = 0.182, km2). This difference is not significant (F = 0.474, p P = 0.699). = 0.504). Factors affecting Home range, Movement and Diet in lions 15

Table 4 Important waterholes with counts and score. Counted time outside the park borders was spent in the cold are the number of lions with a certain waterhole within their dry season (respectively 38% and 22%), less in the home range (95%), core area (50%) or heart of the core area wet season (respectively 8% and 10%) and least in the (5%). Score is calculated by giving waterholes that are situated within 5% kernel 5 points, 50% kernel 3 points and 95% kernel hot dry season (respectively 2% and 6%). 1 point. When a waterhole is situated within a 5% kernel, it also is counted in the 50% kernel and 95% kernel. Only water- Waterholes holes with scores 6 or higher are given. In the cold dry season, no waterholes have a score higher than 5. For most of the lions the heart of the core areas (HM 5% 50% 95% Score 5%), calculated for all data is, located around, or All seasons Gobe 2 2 2 10 close to mares, or in unreachable parts of the park of Touma 0 2 4 8 which the presence of waterholes was unknown (Fig- Magala 0 2 4 8 ure 5Figure 6Figure 7 andFigure 8). As presented in Table 4 most important waterholes in the research pe- Saourware 0 2 3 7 riod turned out to be Gobe (10), Touma (8), Magala Gamzamia 1 1 3 7 (8), Saourware (7) and Gamzamia (7). Gobe, Gamza- Alaza 0 1 4 6 mia and Saourware are waterholes on the floodplain Kouloudika 0 1 4 6 and Touma and Magala are situated within the forest Louloumbaya 1 1 2 6 zone. In the hot dry season, water is only available in Mourgouma 0 1 4 6 few waterholes that contain water permanently. Not Tchikam 0 1 4 6 surprisingly these waterholes are part of the home Hot dry season Gobe 2 2 2 10 ranges of the lions. Vo and Zie are waterholes in the Saourware 0 2 2 6 northeastern part of the floodplain that contain water Talabal 1 1 2 6 permanently, but are no part of lion home ranges. Wet season Touma 0 2 4 8 In the hot dry season, Gobe is for both L2 and L3 part Kouloudika 0 1 4 6 of the heart of the core area (HM 5%). This might Magala 0 1 4 6 imply a social relation between these lions, which Mandoure 0 1 4 6 is confirmed by the fact that they were in the same group when they were collared in may 2007 (Tumen- Time spend outside park border ta, 2007).

When all data of all lions are considered, 13% of the Diet time was spent outside the park borders. Most time was spend outside the park in the cold dry season GPS-clusters were visited for L1, L2, L3 and L5. In (24%), much less in the wet season (8%) and least in total 508 GPS-clusters were selected, 299 were vis- the hot dry season (3%). L4 did not spend any time ited and 143 (48%) carcasses were found (Table 5). outside the park border, but data for her were few and On the 58 random GPS-points we found 19 (33%) data from the cold dry season are absent (Table 3). carcasses. 14 of the random carcasses were very old, 2 old and 3 recent. Differences between the sexes Most important prey species for the four lions for Males (23%) spent considerable more time outside the which the diet reconstruction was done, were kob park borders than females (7%). However this differ- (27%) and cattle (17%). 24% of the total diet of the ence was not signiﬁcant (F = 1.459, p = 0.244). Both four lions consists of livestock. Livestock seems to be males and females spend most time outside the park an important source of food. in the cold dry season (respectively 33% and 28%) Table 5 Numbers and percentages of GPS clusters that were and least in the hot dry season (respectively 6% and selected, visited and where carcasses were found for the four 2%). Remarkable is the difference in the wet season. lions and the total. Males spend 20% outside the park and females 2%. L1 L2 L3 L5 T Selected 161 78 188 81 508 Differences between the habitats clusters Floodplain lions and forest zone lions spent respec- Clusters 84 (52) 41 (53) 117 (62) 56 (69) 298(59) tively 13% and14% outside the park borders. The pat- visited (%) tern of time spent outside the park borders was for Carcasses 41 (49) 30 (73) 46 (39) 26 (46) 143 (48) both ﬂoodplain and woodland lions the same: most found (%) 16 H.D. VISSER ET AL.

Prey consumed by L3 (N=30) Prey consumed by L5 (N=26) Elephant Ostrich Gazelle 3.85% 3.85% Horse 3.33% Reedbuck 6.67% 3.85% Kob Roan Cattle 23.08% 6.67% 3.85% Warthog 7.69% Kob 40.00%

Giraffe Giraffe 20.00% 7.69%

Roan 19.23% Gazelle 11.54%

Topi Cattle 15.38% 23.33%

Prey species consumed by L3 (N=46) Prey consumed by L1 (N=41) Marabou stork Sheep/Goat Jackal Elephant Horse 2.17% 2.17% 2.38% 2.38% 4.35% Monitor lizard Roan 4.76% Cattle 4.35% 21.43% Elephant Giraffe 6.52% Gazelle4.76% Kob 7.14% 34.78%

Sheep/Goat 8.70% Warthog 7.14%

Roan 19.05%

Topi 9.52% Cattle 17.39% Kob Topi 9.52% Giraffe 19.57% 11.90% Figure 9 Preys species consumed percentages by L1, L2, L3 and L4 The four different lions show a difference in species Prey size they eat (Figure 9). The most important prey species for L1 are cattle (21.43%), roan (19.05%) and giraffe For the four lions investigated in this study we found (11.90%). More than 80% of the diet of L2 consists 46% (66) carcasses of large prey and 54% (78) car- of three species: kob (40.00%), cattle (23.33%) and casses of medium prey, which gives a ratio large/me- giraffe (20.00%). L3 mostly eats kob (34.78%), topi dium prey biomass of 0.85. However, in this ratio live- (19.57%) and cattle (17.39%). For L4 kob (23.08%), stock is included, which might give a distorted view. roan (19.23%) and topi (15.38%) are most important When livestock is removed, only 34% of the prey is prey species. large and 66% medium, which gives a large/medium prey biomass ratio of 0.51. Bauer et al.. (2008) found Prey preference a 49% medium sized prey and 51% large sized prey in western and central Africa and 35%Page medium 1 sized Page 1 As presented in Table 6, most preferred species were prey and 65% large sized prey for eastern and south- gazelle (0.96), ostrich (0.91) and roan antelope (0.91). ern Africa. Although livestock is an important part of the diet of some lions, the lions have a negative preference for Differences between the sexes both cattle and sheep/goat. Remarkable is that kob From the prey species that were eaten by the males, and topi, the most abundant species in WNP, are less 56.9% was classiﬁed as large, while females have preferred than other prey species. eaten only 36.1% large prey (Figure 10). When only

Page 1 Page 1 Factors affecting Home range, Movement and Diet in lions 17

100.0% 100.0% Prey size Prey size 80.0% Large 80.0% Large Medium Medium 60.0% 60.0%

40.0% 40.0% Percentage Percentage 20.0% 20.0%

0.0% 0.0% Male Female Floodplain Forest zone Sex Habitat type Figure 10 Prey size for males and females; medium <200 kg, Figure 11 Prey size for lions from the forest zone and wood- large >200 kg. land; medium <200 kg, large >200 kg.

wild prey is considered, males have a higher prefer- dex = 0.01) than for large sized prey (Jacob’s index = ence for large sized prey (Jacob’s index = 0.30) than -0.03), while lions from the forest zone have a higher for medium sized prey (Jacob’s index = -0.22), while preference for large sized prey (Jacob’s index = 0.2) females have a higher preference for medium sized than for medium sized prey (Jacob’s index = -0.11). prey (Jacob’s index = 0.02) than for large sized prey The difference in prey size between lions from the (Jacob’s index = -0.06). Males have an obvious pref- floodplain and the forest zone is not significant (df = erence for large sized prey, but the preferences of fe- 1, p = 0.429). males are close to the ratio of medium sized prey/large prey found in the environment. Males eat significantly Prey type more large prey than females. (df = 1, p = 0.012). For the four lions investigated in this research we Differences between the habitats found 20% livestock and 80% wild prey. Livestock From the prey species that were eaten by lions from the raiding takes mostly place in the cold dry season, 26% floodplain 43.4% was classified as large and 56.6% as of the found carcasses were livestock, less in the wet medium. Lions from the forest zone have eaten 50.0% season, 19% livestock, and least in the hot dry season, large prey and 50.0% medium sized prey. Lions from 14% livestock (Table 3). the floodplain have eaten relatively more large sized prey than lions from the forest. When only wild prey Differences between the sexes is considered, prey preferences for large or medium As presented in Table 3, of the preys eaten by the male sized prey are quite close to 0, which means that rela- lions 25% was livestock and 75% was wild prey. Fe- tive prey consumption is close to relative availability males have eaten 15% livestock and 85% wild prey in the environment. Lions from the floodplain have a (Figure 12). This difference was not significant (df = higher preference for medium sized prey (Jacob’s in- 1, p = 0.146). Males ate most livestock in the wet sea-

L1 L2 L3 L5 Total # % (N=39) JI (N=28) JI (N=43) JI (N=25) JI % JI Elephant 246 1.01 5.13 0.67 0.00 -1.00 2.33 0.40 4.00 0.60 2.96 0.49 Giraffe 604 2.47 12.82 0.68 21.43 0.79 6.98 0.48 8.00 0.53 11.85 0.66 Gazelle 28 0.11 7.69 0.97 3.57 0.94 0.00 -1.00 12.00 0.98 5.19 0.96 Kob 1562 6.38 10.26 0.23 42.86 0.74 37.21 0.71 24.00 0.58 28.15 0.63 Ostrich 9 0.04 0.00 -1.00 0.00 -1.00 0.00 -1.00 4.00 0.98 0.74 0.91 Roan 148 0.60 20.51 0.94 7.14 0.84 4.65 0.77 20.00 0.94 12.59 0.91 antilope Topi 848 3.47 10.26 0.49 0.00 -1.00 20.93 0.72 16.00 0.64 12.59 0.57 Page 1 Page 1 Warthog 21 0.09 7.69 0.98 0.00 -1.00 0.00 -1.00 8.00 0.98 3.70 0.95 Shoats 3543 14.48 2.56 -0.70 0.00 -1.00 9.30 -0.22 0.00 -1.00 3.70 -0.59 Cattle 17459 71.35 23.08 -0.51 25.00 -0.48 18.60 -0.59 4.00 -0.89 18.52 -0.59

Table 6 Prey preference for the four lions assessed as Jacob’s Index (JI). Availability data are coming from an aerial count executed in 2007 (Omondi et al., 2007). Monitor lizard, marabou stork, reedbuck and jackal are excluded from the calculations because no data were available for these species. 18 H.D. VISSER ET AL.

100.0% Prey type 100.0% Prey type 80.0% Livestock 80.0% Livestock Wild prey Wild prey 60.0% 60.0%

40.0% 40.0% Percentage 20.0% Percentage 20.0%

0.0% 0.0% Male Female Floodplain Forest zone Sex Habitat type Figure 12 Prey type for males and females. Figure 13 Prey type for lions from forest zone and ﬂoodplain.

son (28%), less in the cold dry season (23%) and least park and prey diversity (Table 7). Time spent outside in the hot dry season (15%), while females ate most the park was positive correlated and prey diversity livestock in the cold dry season (23%), less in the hot was negatively correlated with livestock predation. dry season (15%) and least in the wet season (4%). Close to significant was the correlation between home range size and livestock predation. This correlation Differences between the habitats was positive. Other relations with livestock predation Lions from the floodplain ate relatively more live- were not significant. stock (30%) than lions from the forest zone (16%) (Figure 13). This difference is not significant (df = 1, DISCUSSION p = 0.124). Lions from the floodplain eat most livestock in the cold dry season (53%), less in the wet Diet season (22%) and least in the hot dry season (10%). Lions from the forest ate most livestock in the hot dry For the lions we investigated we found that they ate season (25%), less in the wet season (15%) and least 34% large sized prey and 66% medium sized prey. in the cold dry season (11%). That results in a large/medium prey ratio of 0.51. Bauer (2008) found a ratio of 0.15, which is way lower. He found that in eastern and southern Africa lions Prey diversity fed on 65% large prey and 35% medium sized prey. The low percentage of large sized prey in the diet is In total 143 prey of 14 different species were found, not a result of low preferences for large prey (large which gives a prey diversity of 0.1. Females had a sized prey: 0.20, medium sized prey: -0.11), but a re- more diverse diet than males (resp. 0.29 and 0.22). sult of low densities of large prey in WNP. The only Lions from the floodplain had a less diverse diet than large animals suitable to hunt for are roan antelope lions from the woodland (resp. 0.20 and 0.30). and juvenile giraffe. Elephants are to big, because of relatively small group sizes (Bauer et al., 2003 ; Kok Factors related to livestock predation et al., 2008). These small group sizes are small because of small prey animals. Significant correlations with seasonal fluctuations in livestock predation were the time spent outside the Cluster method

Table 7 Linear regression calculated for several factors that are In this research we did not assume that carcasses found possibly related with livestock predation; ** = correlation is significant at the 0.01 level, * = correlation is significant at the were actually killed by the research lions, only con- 0.05 level. sumed by them. Van Rijssel (2008), using the same methods and collecting part of the data used in this P R square Pearson Page 1 Page 1 Home range size 0.074 0.345 0.587 study, assumed that all found carcasses on the GPS- Time spent outside park 0.006 0.632 0.795** clusters were killed by the lion under investigation it- Prey diversity 0.044 0.416 -0.645* self. We think that this assumption is wrong for a part of the carcasses. At least a part of the carcasses will Travel distances day 0.815 0.007 -0.085 have another reason of death (i.e. scavenging, animals Travel distances night 0.402 0.089 0.299 died from disease, etc.). As stated before, group sizes Travel distances total 0.406 0.088 0.296 are relatively small in WNP. Animals weighing more Factors affecting Home range, Movement and Diet in lions 19 than 1000 kg are said to be safe from lion predation if possible, to avoid data that are older than one rainy (Schaller, 1972). Nonetheless a substantial part of the season. diet of the lions consisted of adult giraffe and elephant. Another possible limitation to the cluster method It is most unlikely that these animals eaten by lions in is the assumption that lions eat for more than three WNP have been killed by the lions themselves. Dur- hours. This is only true for large prey (Schaller, 1973), ing the fieldwork we observed several fresh elephant small prey species might be overlooked and a bias in- carcasses that died from other causes than lion preda- troduced. A good way to counter this problem is to tion. One sick male elephant was observed in an area identify hears from lion scats and compare the results with obvious lion activity (fresh carcasses of kob and with those of the cluster method. topi were observed on a daily basis). This weak indi- Although the method used in this study has some lim- vidual was alone and got weaker every day. After a itations, it might be the only suitable method. Other few days he died. Two other elephant carcasses were methods are not applicable in areas with small lion killed by poachers and the remains left. The presence population as WNP. Often diet is deduced from direct of adult giraffe and elephant in the diet of the research observations of lions on a kill (Schaller, 1972; Fun- lions underline the fact that a part of the diet is ob- ston, 2001). In WNP, lions are hardly observed while tained by scavenging. When reconstructing the diet eating a prey. Another method is to collect scats and of lions with this cluster method, one cannot deduce identify hears of prey species under a microscope. the time of kill, or draw conclusions about the hunting To link these scats to a specific lion, DNA analysis is behaviour of lions. needed, which makes this method very expensive. During fieldwork we found much carcasses that didn’t belong to the GPS-clusters of the research lions. We Home ranges and movement decided to use a control and searched for random clusters. We found a success score of 33% for these The home ranges of the lions investigated in this random clusters. This was 48% for clusters selected study were large compared to home ranges of lions for the lions. Because of the high presence of random in East and Central Africa (Van Orsdol et al., 1985; carcasses in WNP (and in any other park) an essen- Woodroffe & Ginsberg, 1998). This difference in tial part of the method is the aging of the carcasses. home range size can be explained mainly by the When we found a carcass that differed in age from difference in density of prey species between the the expected age, we did not recognize this carcass as areas (Van Rijssel, 2008). Home range size fluctu- belonging to the lion under investigation. ates strongly over the seasons, with largest home Part of the initially selected GPS-clusters turned out ranges in the wet season, smaller in the cold dry to be too old and we found less carcasses and the car- season and the smallest in the hot dry season. This casses that we found were often too incomplete to pattern is different for daily movement, the lions determine the prey species. During the rainy season moved most during the hot dry season, less dur- carcasses become scattered or disappear into the soil. ing the wet season and the less during the cold dry Our fieldwork showed that GPS-clusters older than season. In the hot dry season lions have small home one rainy season were not reliable anymore. We de- ranges, but move much within this area, while in cided to discard all data that were older than one rainy the cold dry season lions have large home ranges, season. This resulted in a reduction of the selected but move less within this area. The difference be- GPS-clusters for L3 and of all the GPS-clusters of tween movement and home range size might be L4. The success ratio of our search (Table 5) shows explained by the prey type lions hunt. In the hot still poor results for L3 (39%) and maybe even one dry season wild prey is bound to waterholes and rainy season might be too much. The GPS-clusters of present in high densities locally, while in the cold L5 were very recent and the carcasses were easily to dry season wild prey is hardly available and lions identify. Carcasses were quite complete and often fur switch their diet to livestock. As Hemson (2003) was still available. When no carcasses were present, it suggested livestock is a relatively easy prey with was clear that lions had rested at many of these plac- low energy investment to catch. Former studies es: the grass was flattened and scats were present. In have shown and this study has confirmed that when addition, WNP is scattered with carcasses and 74% of lions leave WNP, they feed on livestock (Bauer & these carcasses are classified as very old. When old De Iongh, 2005; Van Bommel et al., 2007; Van Ri- GPS-clusters are investigated, the chance of a wrong jssel et al., 2008;). The lions in this study move less interpretation is considerable higher. It is advisable outside the park than inside the park, which might for further use of this method to use recent data and, be an indication for livestock predation. 20 H.D. VISSER ET AL.

Differences between the sexes ed that the forest zone was a less suitable habitat for lions than the floodplain and that this difference in Between the sexes, we expected that males will suffer quality would be visible in their home ranges, move- more than females from the degradation of WNP. We ments and diet. On the other hand we expected that li- expected that males will have both larger home rang- ons from the floodplain would show more fluctuations es and move more than females, this expectation has over the year. We found some differences, but none been proven significantly true only for the wet season. of them was significant. Lions from the floodplain During the other seasons the females had larger home showed no more fluctuations in movement than lions ranges and moved more than males. However none of from the forest zone and lions from the forest zone did these differences was significant. A remarkable result not move more than lions from the floodplain. Home is the fact that in all seasons males move less than ranges of lions from the forest zone were on average females during daylight. This difference might be ex- larger than lions from the floodplain, but this was no plained by the fact that males have manes with less significant difference. Lions from the floodplain ate efficient body temperature regulation than females as more livestock than lions from the forest zone, but a result (West & Packer, 2002, Patterson et al., 2006). this difference was not significant as well. Remark- To maintain their body temperature, males have to able is not how much the lions from different habitats move less than females during the hot hours of the differ, but how less. A difference in habitat quality day. The less effective body temperature regulation of might exists, but the methods applied in this study did males might explain as well partly the differences be- not detect a difference. tween the home ranges and movements of both males and females. Furthermore we expected that males Factors affecting the lion-livestock conflict contribute more to the lion-livestock conflict than females. This seems true, but the results are not signifi- As we have shown most important factors related to cant. Males ate relatively more livestock in the hot dry livestock predation are time spent outside the park and the wet season, while females ate relatively more borders, prey diversity and home range size. The rela- livestock in the cold dry season. It seems that these tion between time spent outside the park is not sur- findings contradict our expectations. Some aspects of prising, this is only a confirmation of the findings of our fieldwork will affect the findings. During the cold other studies that found livestock raiding mostly out- dry season L1 was part of a group of five lions that side the park borders (Bauer & De Iongh, 2005; Van spent much time on the Cameroon-Nigerian border. Bommel et al., 2007; Van Rijssel, 2008). Only L1 and another lion returned to WNP, the others Two observations are contradicting the predictions were most likely killed. It was not possible to visit we made with the help of optimal foraging theory. this area within this study, but as we have proven in We expected that a. lions will kill livestock mainly in WNP a significant positive correlation exist between difficult times (i.e. cold dry season/wet season) b. in livestock predation and time spent outside park. As difficult times diet will be broader than in easy times in this area hardly any wild prey species live outside and c. in order to find their prey, lions have to travel WNP we can assume that during the time spent in this more each day in times of low prey abundance. Our area, L1 survived on livestock exclusively. The same findings are a negative correlation with opportunism holds for L2. No carcasses were searched for clus- and no correlation with daily travel distance. ters from the cold dry season, while he spent much A possible explanation for this observation is that in time outside the park. For L3 we made an expedition times of low wild prey abundance lions switch their into a hardly penetrable area north of the park. In this diet to an easier to catch prey: livestock. Because live- area we only found carcasses of livestock on the GPS- stock is less preferred than wild prey, this decision is clusters. Male lions did eat significantly more large only taken when times are hard (i.e. when the ratio of prey than female lions. The results of the comparison energy investments in searching s overweigh the total between males and females show that differences be- energy intake). When lions feed on livestock, they between the sexes are of major importance in the ecol- come focus less on hard to catch wild prey (e.g. they ogy of lions and the contribution to the lion-livestock become less opportunistic). In the hot dry season prey conflict. species are bound to waterholes. Lion home ranges are small and waterholes are important elements. In Differences between habitats the wet season when herbivores don’t rely on waterholes anymore, their abundance decreases and lion Between lions from the different habitats, we expect- home ranges become much larger. Because of low Factors affecting Home range, Movement and Diet in lions 21 prey densities and absence of livestock as a replace- important factor. Furthermore we showed that the op- ment, lions have to increase their home range in or- timal foraging theory is a powerful tool in predicting der to find their prey. In the cold dry season three of reactions of lions to fluctuations in ecological factors. the four (no data of L4 were available for this period) Reversibly home range and movement can possibly lions in this research leave the park to feed on live- be used to predict if lions will become problem ani- stock. Because livestock is an relative easy to catch mals. prey, home ranges are smaller in the cold dry season than in the wet season. This line of thought explains Human pressure the fluctuations of the data of both male lions (L1 and L2). L3, a female, shows a different pattern. She has During the fieldwork, much illegal activity was ob- the smallest home range in the hot dry season, larger served. Herdsmen with cattle, poachers or traces of in the wet season and the largest in the cold dry sea- poachers were seen on a daily basis. Campsites with son while she moves most in the hot dry season, less facilities for several people to stay for a longer period in the wet season and the least in the cold dry season. of time were found. Most of these illegal activities Remarkable is the fact that during the wet season she were observed close to waterholes that are impor- stays inside the park and she leaves the park in both tant to lions and other wildlife. People compete with cold dry season and the beginning of the hot dry sea- wildlife for resources. During the fieldwork period of son. During the wet season L3 stays in an impenetra- 2008, students observed lions regularly in the North- ble part of the park and it is not clear what she ate. Eastern floodplain, in the area of Gobe, Gamzamia Obviously she fed on easy to catch prey, since her and Tchikam, while during the fieldwork of 2009 no daily travel distance is relatively low. During the cold lions were observed in this area. It is most likely that dry season she leaves the park both east and north of the disappearance of lions from that part of the flood- the park and feeds mainly on livestock. L1, L2 and L3 plain in WNP is caused by the high human pressure. are all seasonal problem animals. For L4 we did not In this study we found that the North-Eastern part of find any carcasses of livestock, she was no problem the floodplain, including the permanent waterholes animal. The size of her home range shows the pattern Zie and Vo, was no part of lion home ranges. During we predicted with optimal foraging theory with the the fieldwork period of 2009, the North-Eastern part least movement in the hot dry season and the most of the floodplain was heavily overgrazed and not a in the cold dry season. The only discrepancy is that single wild herbivore was observed in this area. movement in the wet season is less than in the hot dry The lion population of WNP is under serious threat season. We don’t know how to explain this. Of L4 we and on the brink of extinction (Tumenta et al., in don’t have sufficient data to draw any conclusions. press; De Iongh et al., 2009). Lions, top predators, For all lions the cold dry season is hardest with largest are major indicators of the quality of the environment. home range and movement for L5, or the highest rate Lions disappear as a result of the degradation of the of livestock predation for L1, L2 and L3. habitat and the disappearance of prey species. 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