The effects of interspecific competition on African cheetah (Acinonyx jubatus) foraging
success and diet breadth
Jamie Smith
November 19, 2012
BIOS 3010
Department of Biological Sciences,
Western Michigan University,
Kalamazoo, Michigan 49008, U.S.A.
Introduction
Interspecific competition is an ecological interaction that can have a powerful
and widespread impact on community composition as species influence each others'
distribution and abundance, potentially determining whether species will be able to coexist
(Henke and Bryant 1999, Begon et al. 2006). Predators also have various levels of impact on
prey populations through top-down effects (Sinclair et al. 2003), although Bertram (1979)
found predation to have a weak effect, if any, on prey populations. Broadly, interspecific
competition results in one of two occurrences: coexistence or competitive exclusion (Begon et
al. 2006), both of which have implications for conservation efforts involving endangered
species (Laurenson 1995, Linnell and Strand 2000). Several large African carnivores are
vulnerable or endangered (IUCN Red List of Threatened Species), and competition for
resources is often a contributing factor to such population reductions (Begon et al. 2006). The
asymmetry of competitive ability among organisms will favor some species while others will
suffer reduced fitness, even if they are able to sustain their populations at reduced levels
(Begon et al. 2006). The intensity of interspecific competition increases with increasing trophic
level as food and energy sources become more limited and thus can be particularly strong
and extensive within large predator guilds (Caro and Stoner 2003).
Interspecific competition has been observed among African predators (Creel
and Creel 1996, Linnell and Strand 2000, Caro and Stoner 2003). Carbone et al. (1999)
estimate that carnivores greater than 20 kg in size typically consume prey that is over 45% of
their own body weight, and many of them hunt similarly sized prey (Owen-Smith and Mills
2007). Prey size also influences feeding frequency; the greater amount consumed, the longer
a predator will wait between feedings. Lions, opting for larger prey, tend to feed about every
three days, while cheetahs, due to their smaller size and hunting style, feed on relatively
smaller prey, but feed more frequently to maintain energy (Schaller 1972). The need to feed
regularly and to a degree that maximizes energy benefits from the costs of stalk-and-chase
hunting, which is common to several large African carnivores, results in a guild of large
predators that form around a particular prey selection (Houston 1979).
The large African predator guild is composed of five carnivore species: the lion
(Panthera leo), leopard (Panthera pardus), cheetah (Acinonyx jubatus), wild dog (Lycaon
pictus) and spotted hyena (Crocuta crocuta) (Bertram 1979, Owen-Smith and Mills 2008).
They interact as competitors for food and space, sharing a similar niche, while still
maintaining some separation through prey specialization (Bertram 1979), and some also act
as predators toward other members of the guild (Caro and Stoner 2003). Caro and Stoner
(2003) found that geographic range overlap between cheetahs and 24 other African predators
exceeded 33 percent, which increases the likelihood of carnivore interactions and potential
competition. Cheetahs cover extensive ground when searching for prey (Caro 1994), and the
majority of their prey is migratory (Bertram 1979), increasing their range and the likelihood of
overlap with other members of the predator guild. As stalk-and-chase hunters that pursue
prey at very high speeds, they tend to hunt more successfully in large, open spaces (Mills et
al. 2004). Prey densities also factor into estimates of space requirements; even in predatorfree environments, if the prey density is low, cheetahs will require more geographical space to
support the population (Lindsey et al. 2011).
Cheetahs tend to hunt medium-sized prey (18-65 kg) (Bloomhall 2001, Hayward
et al. 2006). Lions prefer larger prey than cheetahs, but they frequently take medium-sized
prey, demonstrating a tendency toward opportunistic hunting (Hayward and Kerley 2005),
which creates a dietary overlap between lions and cheetahs despite their differing ideal prey
sizes (Figure 1).
Figure 1. Proportion of kills made by each guild member per body mass of prey. Lions,
hyenas, leopards, and wild dogs all have the potential for dietary overlap with cheetahs
(Owen-Smith and Mills 2007).
Cheetahs are unable to behave as contest competitors because their
populations exist at low densities (Bertram 1979). They tend to be scramble competitors,
rather than interference, probably due to their smaller size relative to other key members of
the guild and their solitary hunting style. Their inability to defend against attacks by lions or
hyena packs makes them subordinate within the guild (Hayward and Slotow 2009) and also
exposes them to a greater risk of contest competition, predominantly through kleptoparasitism
(Caro and Stoner 2003). Cheetahs are among the carnivores most vulnerable to
kleptoparasitism, with as many as 10 species able to act as kleptoparasites (Caro and Stoner
2003). Lions and hyenas are the most common kleptoparasites of cheetah kills, responsible
for 78% and 15% of kleptoparasitism events respectively in one community (Hunter et al.
2007b) (Figure 2) .
Figure 2. The mean time (minutes) of arrival of scavengers after a cheetah begins feeding
(Hunter et al. 2007a).
In addition to competitive pressures, cheetahs are also predated by other
members of the guild. Lions and hyenas are responsible for the majority of cub deaths.
Laurenson (1995) found that approximately 73 percent of cub kills in a Serengeti cheetah
population were caused by lions and 15 percent by hyenas. Predation of adult cheetahs is
less frequent, although it occurs occasionally. This predatory interaction between cheetahs
and their main competitors makes it difficult to distinguish between the relative impacts of
predation and competition on avoidance behavior.
Interspecific competition has ecological significance because it affects
abundance and distribution of species (Begon et al. 2006). Competition between large
carnivores impacts the structure of multiple trophic levels and biodiversity (Terborgh 1992,
Woodroffe and Ginsberg 2005, Terborgh and Estes 2010). My goal is to understand the
effects of interspecific competition on cheetah foraging and diet breadth, so here I review the
literature to test the null hypothesis that interspecific competition has no effect on cheetah diet
breadth and foraging success. My alternate hypothesis is that interspecific competition will
cause cheetahs to have reduced foraging success and limit their diet breadth.
Materials and Methods
I utilized multiple journal databases made available through Western Michigan
University’s library website, primarily Web of Science and ScienceDirect. Approximately
thirteen hours were spent searching databases and browsing the results for relevant articles. I
searched both databases using various combinations of the following search terms: cheetah,
hunt*, forag*, compet*, interspecific, diet, carnivore. ScienceDirect produced greater numbers
of articles per search than Web of Science (Figure 3), but the percentage of relevant articles
retrieved from the search results was much greater with Web of Science (Figure 4).
Figure 3. Number of hits produced per search terms per database. ScienceDirect produced
greater numbers of articles per search than Web of Science.
Figure 4. Percentage of relevant articles found per search terms per database. Web of
Science produced a greater percentage of useful articles per search than ScienceDirect.
Results
Direct Effects of Interspecific Competition
i. Space and resource limitations
Space is among the most limiting resources for African wildlife communities, and
the less space that is available to large carnivores, the more they will be forced to interact.
Lindsey et al. (2011) estimated that a group of ten to fifteen cheetahs requires as much as
466 km2 in predator-free areas, and up to 3,739 km2 when sharing space with competitors. In
spite of their clear need for extensive space, cheetahs have not yet been outcompeted by
stronger competitors. Durant (1998) suggested that cheetahs are able to coexist with superior
competitors because of their mobility and capacity to successfully avoid interacting with
competitors without expending excess energy. Their tendency to forage over broad ranges
and follow migratory herds of prey may help them to avoid aggregations of competitors.
ii. Habitat variation
Cheetahs typically hunt in grasslands, and most studies take place in these
open prairies (Schaller 1972, Durant et al. 1988, Fitzgibbon 1990, Laurenson 1995, Durant
1998). However, a study by Broomhall (2001) has shown that cheetahs can inhabit thickets
and wooded areas, possibly to a) exploit higher prey densities, b) conceal themselves from
prey for stalking, and c) conceal themselves from competitors. They preferred habitats with
both open grasslands and brush/woodland cover, rather than one habitat type exclusively.
Such habitat preference is likely due to the fact that they hunt significantly more often and
make more kills in the savanna, but enjoy reduced kleptoparasitism risk and frequency in the
brush. Greater proportion of cover was associated with shorter chase distance, greater
hunting success, and less kleptoparasitism (Figure 5).
Bissett and Bernard (2006) found similar results, observing that more hunts
occurred in the open than in the thicket, but successful kills were made by both males and
females within the brush. A coalition of three males made two kills out of three attempts in the
brush, as opposed to five kills out of eleven attempts in the open. Females with cubs had
three kills out of five hunts in the thicket, and two kills out of five hunts in the open, while
single females had two unsuccessful hunts in the thicket and three kills out of four hunts in the
open. These results demonstrate that cheetahs, while preferring to hunt in the open, can still
make successful kills in brush. Such a habitat would not be conducive to high speed chases,
but may offer a good trade-off, particularly for females with cubs, by providing concealment
from prey for ease of stalking, from competitors to reduce the risk of kleptoparasitism, and
from predators that might kill the cubs.
Figure 5. Correlation between amount of cover and a) chase distance, b) hunting success,
and c) kleptoparasitism per location (Broomhall 2001).
iii. Avoidance behavior
Laurenson (1995) observed that vigilance at kills was significantly higher among
lactating females and females with cubs than other cheetahs, and concluded that this was
due to the risk of cub predation by other competitors. This behavior was also observed by
Hunter et al. (2007b), suggesting that avoidance behaviors may also be a response to
predation risk. Durant (2000) observed cheetah behavior in response to lion and hyena
playbacks and found that vigilance, movement patterns, and cheetahs were significantly less
likely to hunt after hearing lion playbacks (Figure 6). However, no significant difference in
response depending on sex or reproductive status was observed, implying that avoidance
behavior was a response to competition, rather than risk of predation. Bisset and Bernard's
(2007) study took place in a community with a lion pride, and they observed that only the
coalition would hunt near the lions; every other social group remained at least twice as far
away, presumably to avoid competitive interactions, as females with cubs were
kleptoparasitized three times by lions.
Figure 6. Distance (m) of movement and probability of hunting after dummy playbacks with no
sound (black), hyena playbacks (gray) and lion playbacks (white) (Durant 2000).
Indirect effects of interspecific competition
i. Foraging decisions and hunting success
Cheetahs in the playback experiment that opted to hunt after hearing lion
playbacks were less likely to make kills (Durant 2000). Cooper et al. (2007) found that
foraging decisions were affected by the presence of lions. They did not, however, detect any
significant effect on foraging decisions caused by hyenas. As observed in other studies
(Durant 2000), cheetah behavior implies that they find lions to be more intense competitors
than hyenas.
It is suggested that cheetahs select for low and intermediate density prey
populations because they may seek to avoid competitive interactions with lions and hyenas,
both of which aggregate around high density prey populations (Cooper et al. 2007). However,
it is possible that cheetahs prefer smaller density herds because there is a hunting advantage
(Fitzgibbon 1990).
ii. Diet breadth and prey selection may reduce susceptibility to competition
Cheetahs have fairly high degrees of dietary overlap with their competitors
(Hayward and Kerley 2008) (Figure 1), but they select strongly for medium-sized prey
(Hayward et al. 2006), particularly Thomson's gazelles (Fitzgibbon 1990), and also include
Grant's gazelles, impala, blesbok, and springbok, among others in their diet (Hayward et al.
2006). One specific advantage of hunting relatively smaller prey is the decrease in retention
time after the kill. Medium-sized prey can be consumed at greater speed, reducing the
amount of time spent at the kill, and thus the risk of kleptoparasitism. Medium-sized prey are
also easier for cheetahs to subdue, given that they are solitary hunters and relatively small
predators (Hayward et al. 2006).
iii. Temporal separation between cheetahs and competitors
Cheetahs are most active at dusk, while their most intense competitors, lions
and hyenas, are predominantly nocturnal, creating less than 55 percent activity overlap in
time with the other main guild predators (Hayward and Slotow 2009) (Figure 7). This is a fairly
small overlap relative to the 89 percent overlap suffered by lions and 81 percent by leopards,
and supports Hayward and Slotow's (2009) hypothesis that carnivores subject to intraguild
predation would alternate their activity times with their main competitors to avoid such
interactions. In spite of the dietary overlap between hyenas, lions, and cheetahs, temporal
separation of foraging reduces their competitive interactions (Hanby and Bygott 1979).
Figure 7. Temporal activity overlap among large African predator guild members (Hayward
and Slotow 2009).
Discussion and Conclusion
Based on evidence from my literature review, I reject my null hypothesis that
interspecific competition has no effect on cheetah foraging success and diet breadth, and
accept my alternate hypothesis. Cheetah foraging and diet is influenced by a number of
factors, and the interactions are far more complex than a single cause-and-effect relationship,
but there is evidence supporting the idea that interspecific competition influences foraging
success and diet. Kleptoparasitism directly impacts hunting success, and the risk of
kleptoparasitism can cause avoidance behaviors that impact foraging success and potentially
increase energy costs by causing cheetahs to choose not to hunt, or to hunt less successfully
(Durant 2000).
However, there are a number of ways in which these negative impacts are
balanced, allowing cheetahs to coexist in the community. The flexibility of hunting in wooded
habitats shows potential to increase their overall hunting success by reducing the risk of
kleptoparasitism and providing an environment that increases their stealth (Broomhall 2001,
Bissett and Bernard 2007). Opting to hunt from low-density prey populations reduces the
frequency of their interactions with their main competitors (Cooper et al. 2007), and temporal
separation allows cheetahs to avoid competitive interactions with their nocturnal competitors
(Hayward and Slotow 2009).
Literature Cited
Begon, M., J. L. Harper, and C. R. Townsend. 2006. Ecology : individuals, populations, and
communities. Blackwell Science, Oxford.
Bertram B. C. 1979. Serengeti predators and their social systems. Pages 221-285 In A. R. E.
Sinclair and M. Norton-Griffiths, editors. Serengeti: dynamics of an ecosystem, The
University of Chicago Press, Chicago.Bissett C., R. T. F. Bernard. 2007. Habitat selection
and feeding ecology of the cheetah (Acinonyx jubatus) in thicket vegetation: is the
cheetah a savanna specialist? Journal of Zoology 271:310-317.
Bissett C., and R. T. F. Bernard. 2007. Habitat selection and feeding ecology of the cheetah
(Acinonyx jubatus) in thicket vegetation: is the cheetah a savanna specialist? Journal of
Zoology 271:310-317.
Broomhall L. S. 2001. Cheetah Acinonyx jubatus ecology in the Kruger National Park: a
comparison with other studies across the grassland-woodland gradient in African
savannas. Master's thesis, University of Pretoria.
Carbone C., G. M. Mace, S. C. Roberts, and D. W. Macdonald. 1999. Energetic constraints
on the diet of terrestrial carnivores. Nature 402:286-8.
Caro T. M. 1994. Cheetahs of the Serengeti Plains: Group Living in an Asocial Species.
University of Chicago Press, Illinois, USA.
Caro T. M., and C. J. Stoner. 2003. The potential for interspecific competition among African
carnivores. Biological Conservation 110:67-75.
Cooper A. B., N. Pettorelli, and S. M. Durant. 2007. Large carnivore menus: factors affecting
hunting decisions by cheetahs in the Serengeti. Animal Behaviour 73:651-659.
Creel S., and N. M. Creel. 1996. Limitation of African Wild Dogs by Competition with Larger
Carnivores. Conservation Biology 10:526-538.
Dalerum F., M. J. Somers, K. E. Kunkel, and E. Z. Cameron. 2008. The potential for large
carnivores to act as biodiversity surrogates in southern Africa. Biodiversity and
Conservation 17:2939-2949.
Durant S. M. 2000. Living with the enemy: avoidance of hyenas and lions by cheetahs in the
Serengeti. Behavioral Ecology 11:624-632.
Durant S. M., T. M. Caro, D. A. Collins, R. M. Alawi, and C. D. Fitzgibbon. 1988. Migration
patterns of Thomson's gazelles and cheetahs on the Serengeti Plains. African Journal of
Ecology 26:257-268.
Durant, S. M. 1998. Competition refuges and coexistence: an example from Serengeti
carnivores. Journal of Animal Ecology 67:370-386.
Fitzgibbon C. D. 1990. Mixed-species grouping in Thomson and Grant gazelles - the
antipredator benefits. Animal Behaviour 39:1116-1126.
Hanby J. P., and J. D. Bygott. 1979. Population changes in lions and other predators. Pages
249-262 In A. R. E. Sinclair and M. Norton-Griffiths, editors. Serengeti Dynamics of an
Ecosystem, The University of Chicago Press, Illinois, USA.
Hayward M. W. and R. Slotow. 2009. Temporal partitioning of activity in large african
carnivores: tests of multiple hypotheses. South African Journal of Wildlife Research
39:109-125.
Hayward M. W., M. Hofmeyr, J. O'Brien, and G. I. H. Kerley. 2006. Prey preferences of the
cheetah (Acinonyx jubatus) (Felidae: Carnivora): morphological limitations or the need to
capture rapidly consumable prey before kleptoparasites arrive? Journal of Zoology
270:615-627.
Hayward M. W., and G. I. H. Kerley. 2008. Prey preferences and dietary overlap amongst
Africa's large predators. South African Journal of Wildlife Research 38:93-108.
Hayward M. W., and G. I. H. Kerley. 2005. Prey preferences of the lion (Panthera leo).
Journal of Zoology 267:309-322.
Hayward M. W., J. O'Brien, and G. I. H. Kerley. 2007. Carrying capacity of large African
predators: Predictions and tests. Biological Conservation 139:219-229.
Henke S. E., and F. C. Bryant. 1999. Effects of coyote removal on the faunal community in
western Texas. The Journal of Wildlife Management 63:1066-1081.
Houston D. C. 1979. The adaptations of scavengers. Pages 263-286 In A. R. E. Sinclair and
M. Norton-Griffiths, editors. Serengeti: dynamics of an ecosystem, The University of
Chicago Press, Illinois, USA.
Hunter J. S., S. M. Durant, and T. M. Caro. 2007a. Patterns of scavenger arrival at cheetah
kills in Serengeti National Park Tanzania. African Journal of Ecology 45:275-281.
Hunter J. S., S. M. Durant, and T. M. Caro. 2007b. To flee or not to flee: predator avoidance
by cheetahs at kills. Behavioral Ecology and Sociobiology 61:1033-1042.
IUCN 2012. IUCN Red List of Threatened Species. Version 2012.2. <www.iucnredlist.org>.
Downloaded on 20 November 2012.
Laurenson, M. K. 1995. Implications of high offspring mortality for cheetah population
dynamics. Pages 385-399 In A. R. E. Sinclair, Peter Arcese, editor. Serengeti II:
Dynamics, Management, and Conservation of an Ecosystem edited by University of
Chicago Press, Chicago.
Lindsey P., C. J. Tambling, R. Brummer, H. Davies-Mostert, M. W. Hayward, K. Marnewick,
and D. Parker. 2011. Minimum prey and area requirements of the vulnerable cheetah
Acinonyx jubatus: implications for reintroduction and management of the species in South
Africa. Oryx 45:587-599.
Linnell J. D. C., and O. Strand. 2000. Interference interactions, co-existence and conservation
of mammalian carnivores. Diversity and Distributions 6:169-176.
Mills M., L. S. Broomhall, and J. T. du Toit. 2004. Cheetah Acinonyx jubatus feeding ecology
in the Kruger National Park and a comparison across African savanna habitats: is the
cheetah only a successful hunter on open grassland plains? Wildlife Biology 10:177-186.
Owen-Smith N., and M. G. L. Mills. 2008. Predator-prey size relationships in an African largemammal food web. Journal of Animal Ecology 77:173-183.
Schaller G. B. 1972. The Serengeti lion: a study of predator-prey relations. University of
Chicago Press, Illinois, USA.
Sinclair A. R. E., S. Mduma, and J. S. Brashares. 2003. Patterns of predation in a diverse
predator-prey system. Nature 425:288-90.
Terborgh J. 1992. Maintenance of diversity in tropical forests. Biotropica 24:283-292.
Terborgh J. and J. A. Estes. 2010. Trophic Cascades: Predators, Prey, and the Changing
Dynamics of Nature. Island Press, Washington, DC.
Woodroffe R., J. R. Ginsberg. 2005. King of the beasts? Evidence for guild redundancy
among large mammalian carnivores. Pages 154-175 In J. C. Ray, J. Berger, and K. H.
Redford, editors. Large Carnivores and the Conservation of Biodiversity, Island Press,
Covelo, CA, USA.