ArticlePDF Available

Hunting behavior in West African forest leopards

Authors:

Abstract

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
1,2,
* and Klaus Zuberbu
¨hler
2,3
1
Zoologisches Institut, Universita¨t Bern, Switzerland,
2
Centre Suisse de Recherches Scientifiques, Abidjan, Ivory Coast and
3
School of Psychology,
University of St Andrews, St Andrews, Scotland, UK
Abstract
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
Re
´sume
´
La panthe
`re Panthera pardus est un pre
´dateur majeur de
mammife
`res dans l’e
´cosyste
`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
´tudes
mene
´es dans des habitats ouverts de la savane, tandis que
peu est connu sur les panthe
`res forestie
`res. Les donne
´es
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
panthe
`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
´veloppent
des pre
´fe
´rences de proie fortement individualise
´es. Nos
conclusions de
´fient l’image re
´pandue des panthe
`res comme
pre
´dateurs opportunistes nocturnes.
Introduction
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
¨forest,
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.
Methods
Data were collected by the first author in about 100 km
2
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
2
).
*Correspondence: Suot Aquadotas, 7524 Zuoz, Switzerland.
E-mail: jenny.d@compunet.ch
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)
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
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)
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
Activity (proportion 15-min intervals active)
Time (h since sunrise)
(a)
(b)
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;
Zuberbu
¨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,
1995).
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
s
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
0
30
60
90
120
< 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.
0
10
20
30
40
50
60
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
2
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;
Zuberbu
¨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
range.
References
Bailey, T.N. (1993) The African Leopard. Columbia University Press,
New York.
Bodendorfer, T. (1994) Zur Erna¨hrungsbiologie des Leoparden
Panthera pardus und des Lo¨wen Panthera leo im Comoe
´- und
Marahoue
´National park, Elfenbeinku
¨ste-Eine Untersuchung
anhand von Kotproben. Unpublished Thesis, University of
Vienna, Vienna.
Bothma, J.D. & LeRiche, E.A.N. (1994) Scat analysis and aspects of
defecation in Northern Cape leopard. S. Afr. J. Wildl. Res. 24,
21–25.
Hamilton, P.H. (1976). The Movements of Leopards in Tsavo National
Park, Kenya, as Determined by Radio-Tracking. University of
Nairobi, Nairobi.
Hart, J.A., Katembo,M.&Punga, K. (1996) Diet, prey selection
and ecological relations of leopard and golden cat in the Ituri
forest, Zaire. Afr. J. Ecol. 34, 364–379.
Hoppe-Dominik, B. (1984) Etude du spectre des proies de la pan-
there, Panthera pardus, dans le Parc National de Taı
¨en Co
ˆte
d’Ivoire. Mammalia 48, 477–487.
Jenny, D. (1996) Spatial organization of leopards (Panthera pardus)
in Taı
¨National Park, Ivory Coast: is rain forest habitat a tropical
haven? J. Zool. 240, 427–440.
Johnson, K.G., Wei, W., Reid, D.G. & Jinchu, H. (1993) Food habits
of Asiatic leopards (Panthera pardus) in Wolong Reserve,
Sichuan, China. J. Mammal. 74, 646–650.
Karanth, K.U. & Sunquist, M.E. (1995) Prey selection by tiger,
leopard, and dhole in tropical forests. J. Anim. Ecol. 64,
439–450.
Rabinowitz, A.R. (1989) The density and behavior of large cats in
a dry tropical forest mosaic in Huai Kha Khaeng Wildlife
Sanctuary, Thailand. Nat. Hist. Bull. Siam. Soc. 37, 235–251.
Stuart, C.T. & Stuart, T.D. (1993) Prey of leopards in the western
Soutpansberg, South Africa. J. Afr. Zool. 107, 135–137.
Zuberbu
¨hler,K.&Jenny, D. (2002) Leopard predation and primate
evolution. J. Hum. Evol. 43, 873–886.
Zuberbu
¨hler, K., Jenny,D.&Bshary, R. (1999) The predator
deterrence function of primate alarm calls. Ethology 105,
477–490.
(Manuscript accepted 30 November 2004)
200 David Jenny and Klaus Zuberbu¨hle
2005 African Journal of Ecology, Afr. J. Ecol.,43, 197–200
... Even the same predator species living in two different regions can exhibit contrasting activity patterns, aligning closely with the prey species characteristic of their respective region. This phenomenon, often referred to as the "mirroring effect", has been documented across various predators (Harmsen et al., 2011;Jenny and Zuberbühler, 2005;Kawanishi and Sunquist, 2004). In our study, the snow leopard's peak activity at dawn and dusk coincides with the decreasing activity of the ibex, suggesting a potential 'mirroring Fig. 6. ...
... Allen et al. (2020) observed that leopard distribution varied among the dry and wet seasons, and was primarily affected by interactions with other larger carnivores and vegetation cover, with leopards expanding habitat use in the wet season to areas with less cover. In the wet season, prey abundance is more evenly distributed throughout the landscape (Chaka et al. 2021;Kittle et al. 2017;Patterson et al. 2004), and leopards may use areas of moderate vegetation density more frequently when rainfall provides an additional element of secrecy during hunting (Jenny and Zuberbühler 2005). Conversely, high rainfall negatively influenced detection probability in sites with high vegetation productivity. ...
Article
Full-text available
Maximizing detection probability of elusive species enhances the robustness of population and occupancy estimates, which are essential for supporting impactful conservation strategies. Although the number of camera trap studies on leopards is increasing, few have assessed the drivers influencing leopard detection specifically. We used occupancy modeling embedded in a causal inference framework to provide four biologically robust site use models against which to test the influence of six factors likely influencing leopard detectability at the level of encounter probability, trigger probability, and image quality. In this study, vegetation productivity moderated by rainfall was the top predictor of leopard detectability associated with three of the four site use models. While optimizing detection probability improves estimates of population parameters, the cost‐effectiveness of the study designs is also an essential criterion to consider for long‐term monitoring of elusive species. Camera trap placement involves minimal cost in the early stages of the grid design. Our results suggest that setting cameras in microhabitats of moderate productivity improved leopard detectability in the wet season. This study can inform the design of camera trap studies occurring in semi‐arid bushland ecosystems to improve estimates of leopard population and occupancy.
... They were primarily crepuscular and nocturnal, consistent with earlier findings (Chaudhary et al., 2020;Rafiq et al., 2020) and observed in other felids such as the leopard cat (Austin et al., 2007), cheetah (Rafiq et al., 2020), tiger (Wang et al., 2016;Yang et al., 2018), and lion (Chaudhary et al., 2020). Their heightened activity during twilight and at night is likely due to activity of their prey species (Jenny and Zuberbühler, 2005;Sunquist and Sunquist, 2002). Barking deer and wild boar, its primary prey species in this region (Kunwar and Koju, 2019), are most active during twilight and at night (Fig. 4). ...
... They were primarily crepuscular and nocturnal, consistent with earlier findings (Chaudhary et al., 2020;Rafiq et al., 2020) and observed in other felids such as the leopard cat (Austin et al., 2007), cheetah (Rafiq et al., 2020), tiger (Wang et al., 2016;Yang et al., 2018), and lion (Chaudhary et al., 2020). Their heightened activity during twilight and at night is likely due to activity of their prey species (Jenny and Zuberbühler, 2005;Sunquist and Sunquist, 2002). Barking deer and wild boar, its primary prey species in this region (Kunwar and Koju, 2019), are most active during twilight and at night (Fig. 4). ...
Article
Full-text available
The common leopard (Panthera pardus) is a large carnivore species primarily ranging in the protected areas within the flat plains of Nepal. Facing competition from tigers (Panthera tigris) and decline prey population within these areas, the leopards venture into the mid-hills, leading to increased conflicts with local people. This study was designed to identify the predator and determine the spatiotemporal interactions between predators and potential prey species in the Chhatradev Rural Municipality (CRM) of Arghakhanchi, Nepal. The study utilized 18 camera traps in two phases, with a total sampling effort of 2,402 trap night. Two common leopards and individuals of six prey species were identified. The findings indicated that common leopard activities were primarily nocturnal, with distinct crepuscular peaks at dawn and dusk, when the highest spatiotemporal overlaps were seen with wild boar and barking deer. Composite scores indicated that wild boars and barking deer had relatively higher scores, suggesting that they are the main prey species for common leopards. Our study confirmed the presence of common leopards in the mid-hill regions of Nepal, emphasizing the importance of maintaining a healthy prey population for leopard conservation and human conflict reduction.
... Ultimately, post-fire mediated alterations could affect space use by prey taxa, like small mammals, which are strongly linked to the structure of the vegetation they use as cover (Leahy et al., 2016;Lima and Valone, 1986;Verdolin, 2006;Winnie and Creel, 2017). Furthermore, anti-predatory behaviors of prey could affect predators hunting behavior, triggering a sequence of specific predator-prey interactions (Dawkins and Krebs, 1979;Monterroso et al., 2013;Winnie and Creel, 2017) which may include spatial and temporal adaptations by both prey and predators (Eriksen et al., 2011;Jenny and Zuberbühler, 2005). ...
Article
Full-text available
Effects of fires on carnivores are still poorly understood, particularly in the fire-prone Mediterranean basin. For instance, whether the effects are mainly mediated by prey abundance (Prey Abundance Hypothesis, PAH), prey catchability (Prey Catchability Hypothesis, PCH) or habitat suitability is unclear. Our objective was to investigate the role of fire, mammalian prey abundance, habitat suitability and their interactions in shaping the use of space and time of two carnivores, i.e., the red fox Vulpes vulpes and the stone marten Martes foina. The study area was Monte Pisano (Italy), where 12 km 2 of surface burned in 2018. In early summer 2021 a stratified random sampling design was implemented, with fire and forest type as main strata. Fifty sites were selected, and two infrared cameras were placed at each site. Camera data were used to develop single-species occupancy models for the two predators, whereas time overlap between theme and their prey was evaluated through the Mardia-Watson-Wheeler test. Fox occupancy decreased with increasing herbaceous cover, but only when "mouse and voles" abundance was medium to high, regardless of habitat type. Fox also had significant differences in temporal activity between burnt and unburnt areas, not coupled by a similar pattern for its prey. In contrast, stone marten occupancy mainly depended on canopy cover. The fox could have adapted its hunting strategy to features of the environment and prey abundance, somehow supporting both PCH and PAH. In time, this species could optimize its activity in burnt and unburnt areas according to the brightness of the night. Differences in activity in "mice and voles" were interpreted as anti-predatory responses to the fox. Lastly, the stone marten did not pursue its prey in open areas. In conclusion, carnivores' habitat use and mammalian predator-prey relationships were overall influenced by fire and post-fire successions.
... Leopards living in open environments are known to be largely nocturnal, whereas in forested ecosystems they are more diurnal and crepuscular as they follow activity patterns of their main prey (Jenny and Zuberbühler 2005). This is confirmed by our records for the largely forested Kwano habitat, where some direct sightings occurred during the daytime (cf. ...
Article
Full-text available
West-African populations of the African subspecies of leopards (Panthera pardus pardus) are very fragmented, particularly so in Nigeria, Africa’s most populous nation. We compile historical and recent information about the occurrence of these big cats in what is likely their last remaining stronghold: Gashaka Gumti National Park (GGNP) in Adamawa and Taraba States along the border with Cameroon. Leopards in GGNP’s northern Gumti sector are close to extinction, given that its open plains of savannah are heavily encroached upon by illegal cattle grazing. However, in GGNP’s southern Gashaka sector, leopards are better protected, given its often mountainous and densely forested landscape. We present specific data for a 30 km2 forest-savannah area around a research station at Kwano, which harbours sizeable populations of 35 species of prey species leopards target elsewhere. Here, direct evidence for leopard presence (e.g., footprints, scratchmarks, predation incidences, vocalizations, sightings, cameratrap images) amounts to 1 per year. Albeit this encounter rate is very low, it is relatively steady throughout the last 30 years and into the present. We assume that GGNP’s Gashaka sector represents one of the best chances for leopards to survive in Nigeria.
Article
Characterizing the structure and function of animal communication systems provides insights into the cognitive and evolutionary processes shaping signal complexity. One key question is whether and how call sequences allow potential listeners to make predictions about the call-eliciting referents. Here, we investigated whether primate call sequences contained properties that enabled such predictive processing. We analyzed several years of experimentally elicited alarm responses from a West African forest primate, wild olive colobus monkeys. Using Kullback-Leibler divergence and prediction gain approaches, we identified a simple primate grammar that allowed predictions of referents from only minimal input. In particular, sequence-initial positions reliably discriminated urgent from non-urgent threats while the following positions increased the referential specificity regarding two main predators (eagles and leopards) and non-predatory disturbances (falling tree parts). Sequences often contained further calls, which may allow callers to either confirm the referent or to alter the conveyed information. We concluded that animal communication can contain features adapted for predictive, incremental processing, suggesting evolutionary roots older than language.
Article
Predation risk in the wild varies across the day in a relatively predictable way, as a function of environmental conditions, such as light intensity and temperature, and of predator habits. Prey animals can thus adjust their own activity rhythm to avoid the most dangerous periods. We studied this situation in a coevolutionary perspective, considering that, if prey spread their activity across the day to counter predator temporal preferences, these preferences may in turn change to track prey activity. We therefore built a game-theoretical model to derive evolutionarily stable activity patterns for a predator constantly trying to maximize its chance of capture and a prey trying to minimize this probability. Key parameters concern circadian variations in environmental conditions and their influence on predator hunting efficiency and energy demands of the prey, which dictate its total amount of activity. The model predicts high levels of prey activity during periods of reduced predator efficiency. The predator may then either track these activity peaks and copy the activity patterns of its prey, or concentrate its hunting effort during periods when prey activity is low but conditions favour hunting. In the first case, predator and prey activity patterns will be synchronized. In the second one, they will exhibit strong temporal segregation. We show how these diverging scenarios emerge and how they can help to disentangle the wide variety of situations existing in the wild.
Article
Intraspecific variation in animal movements and its determinants may provide valuable knowledge about ecological and conservation needs of target species. Such studies are relevant to mammalian carnivores due to their wide-ranging behavior and high-profile conservation status. In this large-scale meta-analysis study, I explored the effects of various ecological and anthropogenic factors on daily distances moved by male and female leopards (Panthera pardus) across their global range in Africa and Asia. I tested eight hypotheses by running and ranking 35 linear regression models, separately for each sex. One best male model and two best female models, all statistically significant and having large effect sizes, were built upon 83 cases (45 cases of males and male groups and 38 cases of females and female groups) from 22 publications and 35 study areas. The analysis has shown that movements of male leopards are more intensive in less anthropogenically modified areas, and females move more actively in tropical and subtropical dry forests, shrublands, grasslands and open habitats. Designation and mapping of such areas may allow to delineate possible corridors, identify priority and most sensitive areas for linkages, and plan habitat restoration initiatives for leopard conservation. These results are particularly important for females whose lower mobility and philopatry are arguably among the most limiting factors of population connectivity in wild cats. Additionally, these results have implications for density estimation techniques, reduction of high mortality rates in males, and protection of even less suitable habitats for females.
Article
Full-text available
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.
Article
Full-text available
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.
Article
Full-text available
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.
Article
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
Article
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
Article
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. Résumé On a étudié le régime alimentaire du léopard et du chat doré africain dans la forêt de l'Ituri, au Zaïre, au moyen du contenu des fèces ( N =336) et des carcasses de proies trouvées ( N ‐91). Les félins mangeaient principalement des mammitères (464 des 482 morceaux de proies trouvés dans les fèces). Les ongulés composaient 53,5% et les primates, 25,4% des morceaux de proies identifiés dans les fèces de léopard; on estime d'après les fèces que le poids moyen des proies était de 24,6 kg. Chez le chat doré, 50,9% des fèces identifiées provenaient de rongeurs et 20,2% provenaient d'ongulés; l'estimation du poids moyen des proies était de 1,4 kg. Les léopards étaient à l'origine de 79% des fèces retrouvées et de 74% des traces rapportées lors de comptages systématiques le long des sentiers. Les chats dorés étaient à l'origine de 18% des feces retrouvées et de 15% des comptages de traces. Seize pourcent des fèces de félidés et 11% des comptages de traces n'ont pas pu être attribués avec certitude à l'une ou l'autre espèce. Parmi les espèces de proies de ≥5,0 kg, la prédation des félidés est sélective pour cinq espèces d'ongulés nocturnes et une espèce de primate terrestre diurne. Les primates arboricoles comprenaient 9% des restes des proies dans les fèces. Les proies des aigles couronnés récuperées peuvent être une source importante de primates arboricoles dans le régime alimentaire des félins forestiers. Les chats dorés peuvent être limités par des densités faibles de rongeurs et par la compétition et la prédation par les léopards. Les léopards peuvent être limités par la compétition avec les chasseurs humains pour des proies ongulés dans de nombreuses zones forestières en Afrique.