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Hunting behavior in West African forest leopards



The leopard (Panthera pardus) is a major predator of mammals within the rainforest ecosystem of West Africa. Most of the available information on leopard hunting behaviour comes from studies conducted in open savannah habitats, while little is known about forest leopards. Our radio-tracking data and scat analysis show that forest leopards differ in various ways from the savannah populations. Forest leopards are diurnal and crepuscular hunters who follow the activity pattern of their prey species. They exhibit seasonal differences in activity patterns, and they develop highly individualized prey preferences. These findings challenge the widespread notion of leopards as opportunistic nocturnal predators. La panthère Panthera pardus est un prédateur majeur de mammifères dans l’écosystème de la forêt pluviale en Afrique Occidentale. La plupart des renseignements sur le comportement de chasse des panthères est tirèe d’études menées dans des habitats ouverts de la savane, tandis que peu est connu sur les panthères forestières. Les données que nous avons rassemblées par radio-émetteurs et les analyses des crottes montrent plusieurs différences entre les panthères forestières et des populations de savane. Les panthères forestières sont des chasseurs diurnes et crépusculaires qui suivent le schéma d'activité de leur proie. Ils déploient des différences saisonnières et développent des préférences de proie fortement individualisées. Nos conclusions défient l'image répandue des panthères comme prédateurs opportunistes nocturnes.
Hunting behaviour in West African forest leopards
David Jenny
* and Klaus Zuberbu
Zoologisches Institut, Universita¨t Bern, Switzerland,
Centre Suisse de Recherches Scientifiques, Abidjan, Ivory Coast and
School of Psychology,
University of St Andrews, St Andrews, Scotland, UK
The leopard (Panthera pardus) is a major predator of
mammals within the rainforest ecosystem of West Africa.
Most of the available information on leopard hunting
behaviour comes from studies conducted in open
savannah habitats, while little is known about forest leo-
pards. Our radio-tracking data and scat analysis show that
forest leopards differ in various ways from the savannah
populations. Forest leopards are diurnal and crepuscular
hunters who follow the activity pattern of their prey spe-
cies. They exhibit seasonal differences in activity patterns,
and they develop highly individualized prey preferences.
These findings challenge the widespread notion of leopards
as opportunistic nocturnal predators.
Key words: predation, carnivore, taı
¨forest, selective hunt-
ing, monkey alarm calls
La panthe
`re Panthera pardus est un pre
´dateur majeur de
`res dans l’e
`me de la fore
ˆt pluviale en
Afrique Occidentale. La plupart des renseignements sur le
comportement de chasse des panthe
`res est tire
`e d’e
´es dans des habitats ouverts de la savane, tandis que
peu est connu sur les panthe
`res forestie
`res. Les donne
que nous avons rassemble
´es par radio-e
´metteurs et les
analyses des crottes montrent plusieurs diffe
´rences entre
les panthe
`res forestie
`res et des populations de savane. Les
`res forestie
`res sont des chasseurs diurnes et cre
pusculaires qui suivent le sche
´ma d’activite
´de leur proie.
Ils de
´ploient des diffe
´rences saisonnie
`res et de
des pre
´rences de proie fortement individualise
´es. Nos
conclusions de
´fient l’image re
´pandue des panthe
`res comme
´dateurs opportunistes nocturnes.
It is widely believed that leopards are opportunistic and
nocturnal predators that hunt their prey in proportion to
abundance (Bailey, 1993; Johnson et al., 1993; Stuart &
Stuart, 1993; Bothma & LeRiche, 1994; Bodendorfer,
1994). However, most of the available evidence comes
from savannah habitats while little is known about forest
leopard behaviour. Information from Ituri forest, DR
Congo (Hart, Katembo & Punga, 1996) and Taı
Ivory Coast (Jenny, 1996), suggests that hunting beha-
viour of forest leopards may differ considerably from that
of savannah individuals. In a previous study, we have
documented a large prey spectrum in a population of
forest leopards in the Taı
¨forest (Zuberbu
¨hler & Jenny,
2002). Here, we present data from radio-tracking and
scat analyses to describe the hunting behaviour and in-
dividual prey preferences of forest leopards. Our data
show that forest leopards (a) are diurnally and cre-
puscularly active, (b) exhibit seasonal differences in ac-
tivity, and (c) may develop individual preferences for
particular prey species.
Data were collected by the first author in about 100 km
of undisturbed primary rain forest of Taı
¨National Park,
Ivory Coast (550¢N, 720¢W; Jenny, 1996). Three adult
leopards were captured and equipped with radio-trans-
mitters (Jenny, 1996). Two of them were monitored from
tree platforms during day and night-time intervals
(‘Cosmos’, adult male, 56 kg, 5 February 1993 to 8 May
1994; day: 82 h, night: 23 h; ‘Adele’, adult female,
34 kg, 16 August 1993 to 30 June 1994; day: 244 h,
night: 92 h; night: 18:00–06:00 GMT, corresponding to
the onset of sunset and sunrise). From the platforms it
was possible to score the individuals’ activity as either
‘moving’ or ‘resting’, depending on the stability of the
received signal. Readings were taken every 15 min
(location accuracy ±0.01 km
*Correspondence: Suot Aquadotas, 7524 Zuoz, Switzerland.
2005 African Journal of Ecology, Afr. J. Ecol.,43, 197–200 197
Both individuals were also followed through the forest at
30–150 m (Cosmos, day: 72 h, night: 20 h; Adele, day:
411 h, night: 44 h). Additionally, a third individual was
followed for a short time (‘Cora’; adult female; 32 kg, 14
June 1994 to 28 August 1994; day: 10 h, night: 0 h).
During a proportion of these follows it was also possible
to score activity as either ‘moving’ or ‘resting’ (Adele n ¼
225 h; Cosmos n ¼53.75 h). In addition, it was possible
to make some qualitative observations concerning the
individuals’ hunting behaviour, notably (a) hiding, usually
in dense thickets, (b) approaching monkey groups, and (c)
making kills.
Faecal samples were collected from Adele while follow-
ing her and by tracking her spoor from an infrared-trig-
gered photo-trap set-up along a trail frequently used by
her. This allowed us to compare Adele’s individual prey
spectrum with that of the wider local leopard population
-5 (n = 12)
-4 (n = 12)
-3 (n = 12)
-2 (n = 12)
-1 (n = 12)
Sunrise (n = 12)
1 (n = 11)
2 (n = 16)
3 (n = 29)
4 (n = 43)
5 (n = 58)
6 (n = 50)
7 (n = 56)
8 (n = 55)
9 (n = 50)
10 (n = 47)
11 (n = 34)
12 (n = 17)
Sunset (n = 20)
-10 (n = 20)
-9 (n = 17)
-8 (n = 16)
-7 (n = 12)
-6 (n = 12)
Activity (proportion 15-min intervals active)
Time (h since sunrise)
Cosmos (n = 158.75 h)
Adele (n = 561 h)
-5 (n = 48)
-4 (n = 48)
-3 (n = 48)
-2 (n = 48)
-1 (n = 48)
Sunrise (n = 48)
1 (n = 47)
2 (n = 76)
3 (n = 116)
4 (n = 165)
5 (n = 207)
6 (n = 205)
7 (n = 223)
8 (n = 213)
9 (n = 178)
10 (n = 144)
11 (n = 62)
12 (n = 35)
Sunset (n = 44)
-10 (n = 48)
-9 (n = 46)
-8 (n = 51)
-7 (n = 48)
-6 (n = 48)
Activity (proportion 15-min intervals active)
Time (h since sunrise)
Fig 1 Relative activity patterns of two
radio-collared leopards plotted as a func-
tion of onset of sunrise and sunset (data
from platform monitoring and direct fol-
lows combined). (a) Cosmos (n ¼
158.75 h); (b) Adele (n ¼561 h).
198 David Jenny and Klaus Zuberbu¨hle
2005 African Journal of Ecology, Afr. J. Ecol.,43, 197–200
as determined by scat surveys (Hoppe-Dominik, 1984;
¨hler & Jenny, 2002).
Results and discussion
Activity patterns
Platform monitoring indicated that both individuals were
more active during the day (46.5%) than at night (28.9%),
in sharp contrast to savannah individuals (Hamilton,
1976; Bailey, 1993). Relative peaks at dawn and dusk
corresponded closely with sunrise and sunset (Fig. 1).
Direct follows of leopards at night are extremely difficult to
conduct in dense rainforest habitat. In total, we managed
to obtain ranging data for n ¼179 h combined over the
16-month study period (platform data and focal follows).
These data revealed that at night activity patterns typically
showed only one of two patterns: (a) complete inactivity
(>10 h) or (b) travelling over large distances, suggesting
that although our data set is small, it accurately represents
the nocturnal activity patterns. Daytime activity showed a
more evenly distributed pattern, but inactive periods were
always short (<5 h). This activity pattern is comparable
with that of Asian forest leopards (Karanth & Sunquist,
For Adele, the lowest monthly activity rates were
observed during the rainy period, perhaps because heavy
rainfall increased her hunting success. Percent activity per
month was significantly negatively correlated with rainfall
(Spearman-rank correlation, n ¼11, r
¼)0.718, z ¼
)2.271, P< 0.03).
Ad libitum observations during direct follows revealed
two interesting aspects of hunting behaviour: (a) during
periods of inactivity the three individuals often hide in
dense thickets and (b) after making a kill, they remained at
the same place for two to three consecutive days.
Similar to savannah individuals there was a distinct
sex-difference in ranging behaviour (Rabinowitz, 1989).
Cosmos was moving in 40.4% of all monitored intervals
(n ¼53.75 h), while Adele only moved during 20.1% of
time (n ¼225 h). Adele tended to remain within a relat-
ively small core area of about one hectare (14.2% of
intervals; n ¼561 h; platform and focal data combined).
During daytime follows, a monkey group came within
50 m of the hiding leopard in 60 of 97 hiding bouts (7.4%
of time; n ¼130 h). In contrast, when the observer sat at
one of ten randomly chosen observation points throughout
the study area, monkeys came within 50 m only four of
ten times (1.9% of time; n ¼99 h; Fig. 2), a statistically
significant difference (z ¼)3.092; P< 0.01; Binomial
test; two-tailed), suggesting that leopards selectively chose
hiding spots close to monkey groups (Zuberbu
¨hler, Jenny &
Bshary, 1999).
Prey spectrum
Adele was not an opportunistic hunter but developed in-
dividual prey preferences. First, Adele avoided chimpanzee
parties: in six of fifteen cases of drumming and/or
screaming of a chimpanzee party 100–250 m away, she
< 50 m > 50 m
Distance to next monkey group
Observation hours
With leopard
Without leopard
Fig 2 Number of observation hours the observer was sitting close
(<50 m) or far (>50 m) from the nearest monkey group after (a)
having followed Adele (black bars) or (b) having walked to one of
ten randomly selected points throughout the study area.
Duikers Monkeys Pangolins Other prey
Percent of prey spectrum
Adele (n = 38)
Others (n = 162)
Fig 3 Prey spectrum of focal animal Adele compared with the
average prey spectrum of other leopards in the study area. Duikers
and monkeys all weigh several kilograms and are amongst the
biggest mammalian prey available in the forest.
Hunting behaviour in forest leopards 199
2005 African Journal of Ecology, Afr. J. Ecol.,43, 197–200
started moving in the opposite direction or changed
direction when already moving. Approach was never
recorded. However, leopards may scavenge on already
dead chimpanzees: In two cases, a female or a subadult
male leopard dragged a dead juvenile chimpanzee (10–
20 kg) for 50–80 m before devouring it. Second, Adele
consumed duikers and monkeys significantly more often
than other leopards (v
¼41.49, d.f. ¼8, P< 0.01;
Fig. 3). A successful attack on a primate was observed once
during Adele’s 91 observed hiding bouts when she killed a
sooty mangabey, Cercocebus atys. Third, Adele rarely con-
sumed pangolins, although pangolin remains were com-
mon in other leopard faeces (Hoppe-Dominik, 1984;
¨hler & Jenny, 2002). Thirty-eight fresh faeces
could be assigned to Adele with reasonable confidence.
Only one sample (2.6%; n ¼38) contained pangolin scales
(population average: 26.7%; n ¼150; Fig. 3). Moreover,
leopard faeces containing pangolin remains varied both
regionally and temporarily. In the south-east part of the
study area, 40.0% of faeces (n ¼50) contained pangolin
remains, whereas in the north-west this was only true for
15.2% of faeces (n ¼138). In addition, in the north-west a
sudden and dramatic change in faeces containing pangolin
remains coincided with the suspected death of a resident
leopard: Before March 1993, 32.7% (n ¼55) of faeces
contained pangolin remains. Afterwards, the rate dropped
to 3.6% (n ¼83). No comparable change was observed in
the south-east (before March: 36.7%; n ¼30;after March:
45.0%; n ¼20). There were no obvious habitat differences
across the study area that could have explained uneven
distribution of prey animals.
In sum, our data suggest that forest leopards may differ
from savannah individuals in important ways. First, they
are predominantly diurnally and crepuscularly active.
Second, one focal animal did not consume some potential
prey species, such as chimpanzees and pangolins, but often
others, such as duikers and monkeys. One possible inter-
pretation of this finding is that forest leopards develop
individual prey preferences. The population density of
leopards is high in the Taı
¨forest with substantial range
overlap particularly between the sexes (Jenny, 1996),
suggesting that individuals experience strong competition
forcing them to use different resources within the shared
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2005 African Journal of Ecology, Afr. J. Ecol.,43, 197–200
... This trade-off is dependent on the predators' hunting mode, landscape cover and prey behaviour [15][16][17] . For example, stalk and ambush predators, which largely depend on some form of concealment to approach their prey 14,16 , may use darkness as a substitute for physical cover 18 and shift from diurnal to nocturnal hunting as habitats vary from closed to open, despite the main activity of their prey being diurnal [19][20][21] . Under such conditions, the effectiveness of darkness for concealment may decrease in relation to moon luminosity 18,22 , with predator hunting success varying with the moon phase 23 . ...
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Background Large-scale changes in habitat conditions due to human modifications and climate change require management practices to consider how species communities can alter amidst these changes. Understanding species interactions across the gradient of space, anthropogenic pressure, and season provide the opportunity to anticipate possible dynamics in the changing scenarios. We studied the interspecific interactions of carnivore species in a high-altitude ecosystem over seasonal (summer and winter) and resource gradients (livestock grazing) to assess the impact of changing abiotic and biotic settings on coexistence. Methods The study was conducted in the Upper Bhagirathi basin, Western Himalaya, India. We analyzed around 4 years of camera trap monitoring data to understand seasonal spatial and temporal interactions of the snow leopard with common leopard and woolly wolf were assessed in the greater and trans-Himalayan habitats, respectively. We used two species occupancy models to assess spatial interactions, and circadian activity patterns were used to assess seasonal temporal overlap amongst carnivores. In addition, we examined scats to understand the commonalities in prey selection. Results The result showed that although snow leopard and wolves depend on the same limited prey species and show high temporal overlap, habitat heterogeneity and differential habitat use facilitate co-occurrence between these two predators. Snow leopard and common leopard were spatially independent in the summer. Conversely, the common leopard negatively influences the space use of snow leopard in the winter. Limited prey resources (lack of livestock), restricted space (due to snow cover), and similar activity patterns in winter might result in strong competition, causing these species to avoid each other on a spatial scale. The study showed that in addition to species traits and size, ecological settings also play a significant role in deciding the intensity of competition between large carnivores. Climate change and habitat shifts are predicted to increase the spatial overlap between snow leopard and co-predators in the future. In such scenarios, wolves and snow leopards may coexist in a topographically diverse environment, provided sufficient prey are available. However, shifts in tree line might lead to severe competition between common leopards and snow leopards, which could be detrimental to the latter. Further monitoring of resource use across abiotic and biotic environments may improve our understanding of how changing ecological conditions can affect resource partitioning between snow leopards and predators.
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Primates of modern aspect (euprimates) are characterized by a suite of characteristics (e.g., convergent orbits, grasping hands and feet, reduced claws, and leaping), but the selective pressures responsible for the evolution of these euprimate characteristics have long remained controversial. Here, we used a molecular phyloecological approach to determine the diet of the common ancestor of living primates (CALP), and the results showed that the CALP had increased carnivory. Given the carnivory of the CALP, along with the general observation that orbital convergence is largely restricted to ambush predators, our study suggests that the euprimate characteristics could have been more specifically adapted for ambush predation. In particular, our behavior experiment further shows that nonclaw climbing can significantly reduce noises, which could benefit the ancestral euprimates' stalking to ambush their prey in trees. Therefore, our study suggests that the distinctive euprimate characteristics may have evolved as their specialized adaptation for ambush predation in arboreal environments.
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Human–wildlife conflict is increasing globally, particularly in the areas, where wildlife and humans coexist and share resources. Large mammalian predators such as common leopards not only kill livestock but they are also killing humans. Baitadi is among the top ranked districts in Nepal in terms of number of human common leopard conflict events in last 10 years. The fieldwork for this study was carried out between January and June 2020 in the villages of Bishalpur, Udayadev, Pancheshor and Aamchaura of Baitadi district. Field observation, questionnaire survey, key informant interview and literature review were used for the data collection. Our study found that common leopards killed 23 and injured eight people between 2011 and 2019 in the district. In retaliation, people killed 26 common leopards in the same period, which must have spelt disaster for these rare cats. Despite the increasing number of conflict events, the local people, in general, were found to have positive attitude towards wildlife conservation. Therefore, improved prey species management, awareness raising among the local people and detailed study on habitat assessment, population status of leopards and their prey species are the urgent needs for the mitigation of human common leopard conflict in the district.
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Analysis of feces showed Asiatic leopards (Panthera pardus fusea) to consume a varied diet over a 7-year period. Tufted deer (Elaphodus cephalophus) was replaced as the most frequent prey by bamboo rats (Rhizomys sinense). Fifteen species of large and medium-sized mammals composed the majority of the diet with pheasants, livestock, grass, and soil being eaten occasionally. Reasons for dietary shifts were unclear and may reflect leopards hunting for any readily captured prey by opportunistic encounter and perhaps by changes in hunting behavior, prey availability, or prey vulnerability associated with a bamboo die-off.
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It is generally assumed that alarm calls function in intraspecific communication, for example to warn close relatives about the presence of a predator. However, an alternative hypothesis suggests that, in some cases, signallers may also gain fitness benefits in directly communicating to the predator, for example by advertising perception and unprofitability to predators that depend on unprepared prey. In this study, we show that six monkey species in Taï forest, Ivory Coast, produce significantly more alarm calls to leopards than to chimpanzees, although both are notorious monkey predators. The conspicuously high vocalization rates to leopards had adaptive consequences for the monkeys. By following a radio-collared leopard, we found that after detection and high alarm call rates the leopard gave up its hiding location and left the group significantly faster than would be expected by chance. We discuss these data with respect to the various functional hypothesis of alarm call behaviour and conclude that the high alarm call rates to leopards are part of an anti-predator strategy in primates that may have evolved to deter predators that depend on surprise.
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Between June 1992 and July 1994, two female leopards and one male were radio-tracked. Regular locations of the leopards, the use of a phototrap, and spoor data, provided the first detailed ecological data about this elusive felid in tropical rainforest habitat. The home range of the male was 86 km2, those of the two females were 29 km2 and 22 km2, respectively. One female's home range was fully included within that of the male. Home ranges of neighbouring residents were not exclusive. Population density is estimated at one leopard per 9-14 km2. Intraspecific interactions were rare and predominantly involved mating. The large size of the home ranges and a relatively high population density imply large overlap between adjacent resident leopards' ranges. Differences in the leopard's land tenure system between the rainforest and the savanna are discussed. Doubt is cast on the validity of the often-quoted estimate of one leopard per 1 km2 in tropical rainforest habitat.
Analysis of faecal samples and observations of leopard kills show a total of 32 species of prey. A breakdown of prey species show 99% mammals with a predominance of bovids, followed by monkeys and rodents. The leopard is an opportunist, preying on any species that is available. -from English summary
1. Ecological factors influencing prey selection by tiger Panthera tigris, leopard Panthera pardus and dhole Cuon alpinus were investigated in an intact assemblage of large mammals in the tropical forests of Nagarahole, southern India, between 1986 and 1990. 2. Densities of large herbivores were estimated using line transects, and population structures from area counts. Carnivore diets were determined from analyses of scats (faeces) and kills. Selectivity for prey species was inferred from likelihood ratio tests comparing observed counts of scats to hypothesized scat frequencies generated from prey density estimates using parametric bootstrap simulations. Predator selectivity for size, age, sex and physical condition of prey was estimated using selection indices. 3. Ungulate and primate prey attained a density of 91 animals km-2 and comprised 89-98% of the biomass killed. Predators showed significant (P
Diets of leopard and African golden cat were studied in the Ituri Forest of Zaire by means of scat contents (N=336) and prey carcass finds (N=91). Felids consumed predominantly mammalian prey (464 of 482 prey items in scats). Ungulates comprised 53·5% and primates 25·4% of prey items identified in leopard scats; mean prey weight estimated from scats was 24·6 kg. In golden cat scats 50·9% of identified prey items were rodents and 20·2% were ungulates; mean prey weight estimated from scats was 1·4 kg. Leopards accounted for 79% of recovered scats and for 74% of tracks recorded on systematic counts along paths. Golden cats accounted for 18% of recovered scats and for 15% of track counts. Sixteen percent of felid scats recovered and 11% of track counts could not be attributed to one species or the other with certainty. Among prey species ≥5·0 kg, felid predation is selective for five species of nocturnal ungulates and one species of diurnal, terrestrial primate. Arboreal primates comprised 9% of prey items in scats. Scavenged crowned eagle kills may be an important source of arboreal primates in forest felid diets. Golden cats may be limited by low rodent densities and by competition with and predation by leopards. Leopards may be limited by competition with human hunters for ungulate prey in many areas of forested Africa.