NEW!
learn about the ecological crisis and its solution
click "anthology" on menu below

Haemig PD  (2007 Sympatric Tiger and Leopard.  ECOLOGY.INFO 22

Sympatric Tiger and Leopard

Note: This online review is updated and revised continuously, as soon as results of new scientific research become available.  It therefore presents state-of-the-art information on the topic it covers.

Throughout much of its range in Asia, the tiger (Panthera tigris) is sympatric with the leopard (Panthera pardus).  Because both cats are large in size and eat primarily ungulates (hoofed mammals), ecologists wonder how they can coexist in the same areas without one being more successful than the other and eventually replacing it.  In this review, we compare the habits of sympatric tiger and leopard and summarize the important differences between them that allow coexistence.

To fully understand coexistence, however, we must also look at other predators that compete with the tiger and leopard.  For example, the dhole (Cuon alpinus), a relative of the wolf (Canis lupus), is found in many of the same localities as the tiger and leopard and eats the same ungulates.  Like them, it also actively searches for its prey.  Although smaller in size than the two cats, the dhole is an especially formidable predator because it hunts in groups.  Since the tiger, leopard and dhole often live in the same areas and eat many of the same prey, we will expand this review to examine how all three predators differentiate themselves from each other in order to avoid resource competition and violent encounters.

Differences in Body Size and Prey Size

The tiger averages four times heavier in weight than the leopard (Seidensticker 1976).  The dhole is even smaller than the leopard.

The immense size of the tiger means that it is able to subdue much larger prey than the leopard or dhole.  For example, in the tropical forests of Nagarahole National Park, southern India, where ungulates were very abundant, Karanth and Sunquist (1995) found that tiger usually selected prey weighing more than 176 kilograms, while leopard and dhole "focused on prey in the 31-175 kilogram size class."

Predation on Elephants and Rhinos

Adult elephants and rhinoceroses are so large that the tiger, leopard and dhole usually do not attack them.  However, some exceptions occur.  The World Wildlife Fund in Nepal is currently taking care of an orphaned Indian Rhinoceros (Rhinoceros unicornis) whose mother was killed by a tiger.

Tigers prey more frequently upon elephant calves and rhino calves.  In the tropical forests of Nagarahole National Park, Karanth and Sunquist (1995) found that one percent of all tiger kills were calves of the Asian elephant (Elephas maximus).  Dhole and leopard in the same area did not prey upon elephant calves.

In Chitawan National Park, Nepal, Seidensticker (1976) recorded tiger predation on a calf of the Indian rhinoceros.  In contrast, the leopard did not prey upon rhino calves. Predation by dhole was not studied.

 

Predation on Hoofed Mammals

In the tropical forests of Nagarahole National Park, Karanth and Sunquist (1995) found that five species of ungulates (hoofed mammals) comprised 89-98% of the diet of sympatric tiger, leopard and dhole.  The five ungulates were gaur (Bos gaurus), chital (Axis axis), sambar (Cervus unicolor), muntjac (Muntiacus muntjak) and wild pig (Sus scrofa).  All three carnivores preyed on each ungulate species, but there were some differences in frequency.  Tiger preferred gaur and avoided muntjac, while the leopard avoided wild pig.  Dhole predation on these ungulates did not differ from random.

At the Bandipur Tiger Reserve, southern India, tiger preferred sambar and avoided chital (Johnsingh 1983).  One explanation was that chital "assemble in open areas to spend the night where they are immune to tiger predation (Johnsingh 1983)."  Also, sambar spend more time in the dense vegetation where tigers prefer to make kills, live in smaller groups and do not rest in open areas (Johnsingh 1983).  Leopard did not kill wild pig, and dhole rarely killed it, but tiger readily killed large adult boars, which live alone (Johnsingh 1983).

Predation on Different Sexes and Age Groups of Ungulates

At Nagarahole, Karanth and Sunquist (1995) compared predation on the various sexes of each ungulate species.  Tiger killed more adult males than adult females of the following species: chital, sambar and wild pig.  Dholes also preyed more upon adult male than adult female chital.  These researchers explained the greater predation on male sambar and wild pig by the fact that males of these species tend to live alone and so probably do not detect predators as quickly as females, who live in groups.  They likewise hypothesized that male chital may be more vulnerable to predation than females because of their spacing behavior in large groups. 

At Bandipur, dhole preferred male sambar to female sambar (Johnsingh 1983).  Dhole also preferred to kill chital males that had longer antlers.  One possible explanation for this latter finding is that "stags with large antlers may be hampered when running through dense vegetation and are easily killed (Johnsingh 1983)."

At Nagarahole, tiger predation on gaur was biased towards young gaur, suggesting that the tiger preferentially targets this age-class of gaur (Karanth and Sunquist 1995).

Predation on Monkeys

The hanuman langur (Presbytis entellus) is a leaf-eating monkey.  In the tropical forests of Nagarahole National Park, hanuman langur remains were found in 38% of leopard scats, 19% of tiger scats and only 1% of dhole scats (Karanth and Sunquist 1995).   Furthurmore, langur was under-represented in the diets of both the tiger and dhole, indicating avoidance (Karanth and Sunquist 1995).

The more frequent predation by leopard on langur can be explained by the fact that the leopard climbs trees better than the tiger (Karanth and Sunquist 1995).  The dhole does not climb trees, and its hunting style of running openly in packs may mean that langurs can detect it easier than tiger or leopard, which stalk and ambush their prey (Karanth and Sunquist 1995).

A conflicting result was reported from the Sariska Tiger Reserve, Rajasthan, India, where langur was found more frequently in tiger scats (16.4%) than in leopard scats (6.4%) (Sankar and Johnsingh 2002).  The reason for this reversal with the results from Nagarahole is unknown and worthy of further investigation.

Predation on Smaller Prey

At the Sariska Tiger Reserve, Sankar and Johnsingh (2002) found that rodents (mainly Indian gerbile Tatera indica) and insectivores (mainly grey musk shrew Suncus murinus) were found in 45% of leopard scats, but in only 5% if tiger scats.  They believed that at the time of their study, there was a high availability of rodents and insectivores at Sariska.

At Nagarahole, both leopard and dhole fed more frequently on black-naped hare (Lepus nigricollis) and porcupine (Hystrix indica) than did tiger (Karanth and Sunquist 1995).

 

Food Caching

The leopard differs from the tiger and dhole in that it often takes the animals it kills up into the trees to eat and cache for future consumption.  Here the carcass is usually safe from tiger and dhole, as well as from many other scavengers.  In Chitawan National Park, Nepal, Seidensticker (1976) found that "leopards pulled about half of their kills into trees." 

Both the tiger and leopard hide their kills in dense cover, while dhole leave most of their kills in the open (Karanth and Sunquist 2000).  Gaur carcasses, however, which are too heavy to be dragged, are often left in the open by tiger (Karanth and Sunquist 2000).

Activity Periods

Although the tiger, leopard and dhole hunt round the clock, the dhole differs from the cats by being mainly diurnal (Johnsingh 1983; Venkataraman et al. 1995; Karanth and Sunquist 2000).  At Nagarahole, for example, dhole killed prey most often in the morning (62%) and afternoon (17%), while tiger and leopard killed prey most often during night, evening and morning (Karanth and Sunquist 2000).  The dhole was the only one of the 3 predators that killed prey frequently in the afternoon (Karanth and Sunquist 2000). 

In Chitawan National Park, both the tiger and leopard were found to be "mainly nocturnal," but leopards seemed to "move less often and spend more time in each spot.  Tigers also employed the move-and-stop hunting technique but never stayed as long as leopards in any one place (Sunquist and Sunquist 2002, p. 107)."

Microhabitat Use

At Nagarahole, tiger attacked 81% of their prey in "dense or moderate cover (Karanth and Sunquist 2000)."   An exception was when tiger hunted gaur.  At this time, they usually attacked in more open cover, possibly because gaur are so dangerous (Karanth and Sunquist 2000). 

Leopards at Nagarahole attacked 41% of their prey in the open, twice the frequency of tiger (19%); 28% of leopard kills were in short grass clearings compared to only 8% of tiger kills  (Karanth and Sunquist 2000).  Similar results were found at Bandipur by Johnsingh (1983).  Here, all three predators made more kills in dense cover, but the dhole and leopard also made many of their kills in open places, while the tiger rarely did.

Because the leopard is much smaller than the tiger, it may be able to escape detection better than the tiger while stalking prey in more open habitats (Karanth and Sunquist 2000).  Johnsingh (1983) pointed out that, in general, large cats like the lion (Panthera leo) and tiger "rarely kill prey on short grass or open habitats." 

Although the tiger does not kill often in open habitats, it frequently kills on the edges of them.  At Nagarahole, for example, 45% of tiger kills occurred less than 25 meters from short-grass clearings (Karanth and Sunquist 2000).

Response to Burning of Grassland

In Chitawan National Park, a radio-tracked female tiger and a radio-tracked leopard differed in their responses to the burning of grassland, which transformed areas of dense cover into areas of marginal cover (Seidensticker 1976).  Leopard used the burned areas more frequently than tiger. "Leopard appeared to make direct movements to burned areas immediately after the fire," while tiger rarely did so (Seidensticker 1976)."  Hog deer (Axis porcinus) remained in the burned grassland, especially in small unburned patches, and were hunted by leopard (Seidensticker 1976; Sunquist and Sunquist 2002, p. 106).

A short time after the fire, new grass shoots began to grow and these "attracted large herds of chital and hog deer," but there was not enough stalking cover for the tiger, which continued to hunt in the forest (Sunquist and Sunquist 2002. p. 106).

One month after fire, the grass had grown back to a height of one meter, enough to provide stalking cover for tigers.  At this time, the tiger began to visit the area again on a frequent basis, and the leopard shifted to the forest  (Seidensticker 1976; Sunquist and Sunquist 2002, p. 106).

Utilization of Roads

In Chitawan National Park, tigers frequently walked along roads and trails, while leopards did so only infrequently (Seidensticker 1976; Sunquist and Sunquist 2002, p. 106)

 

Intraguild Predation and Avoidance

Intraguild predation is a term used by ecologists to describe the situation where predators with similar (overlapping) food habits prey on each other.  There may be several advantages to doing this, including direct removal of competitors and intimidation of surviving individuals of the victim species.  The latter may move to habitats where the killer species is absent, change the time of day they hunt so as to reduce encounters with the killer species, or aggregate into groups that can repel the killer species (Palomares and Caro 1999). 

Several studies report intraguild predation between some of the predators reviewed in this article.  Tiger predation on leopard is reported from Chitawan National Park, (Seidensticker 1976), and the Bandipur Tiger Reserve (Johnsingh 1979, 1992).  Tiger predation on dhole is reported from Nagarahole National Park, where a tiger killed 2 dholes while taking over the carcass of an animal they had killed (K.M. Chinnappa in Karanth and Sunquist 2000).  Leopard predation on dhole is reported from both Bandipur (Johnsingh 1983, 1992) and Nagarahole (Karanth and Sunquist 1995).

At Royal Bardia National Park, Nepal, Støen and Wegge (1996) found that tigers occupied the center of the park, while leopards "appeared to avoid the inner areas of the park and were probably confined to the edges and buffer zones between the park and village areas."  These researchers postulated that predation by tiger on leopard forced the latter to avoid the inner areas of the park preferred by tiger.  An indirect result of this displacement of leopards by tiger was that leopards killed many more domestic livestock outside the park than tiger.  Schaller (1967) reported a similar situation at Kanha National Park, central India.

Killing of leopards by tigers thus appears to have important economic consequences in some regions.  However, such intraguild predation has not been well studied and is not well-understood.  In some areas, like Nagarahole National Park, there is complete overlap in areas used by tiger, leopard and dhole, and the type of spatial segregation reported at Royal Bardia National Park does not occur (Karanth and Sunquist 2000).  Nagarhole has an abundance of both large and medium-sized ungulates, providing adequate food for both tiger and leopard, while large ungulates are scarce at Royal Bardia, causing tiger to shift to the smaller ungulates preferred by leopard (Støen and Wegge 1996; Karanth and Sunquist 2000).

Sunquist and Sunquist (2002, p. 107) conclude that the leopard is able to coexist with the larger, socially dominant tiger, "mainly by avoiding hunting places and rest sites tigers prefer. To do this, leopards need a supply of smaller prey and a vegetation type that enables them to avoid tigers."

Editor's Note:  ECOLOGY.INFO also publishes a poem about the tiger and the crisis it faces as an endangered species.  To read this poem, click the following link:  Tiger Tiger Revisited

 

References

Johnsingh AJT  (1979)  Evidence for a tiger eating a panther cub.  Journal of the Bombay Natural History Society 76: 152-153

Johnsingh AJT  (1983)  Large mammalian prey-predators in Bandipur.  Journal of the Bombay Natural History Society 80: 1-57

Johnsingh AJT  (1992)  Prey selection in three sympatric carnivores in Bandipur.  Mammalia 56: 517-526

Karanth KU, Sunquist ME  (1995)  Prey selection by tiger, leopard and dhole in tropical forests.  Journal of Animal Ecology 64: 439-450

Karanth KU, Sunquist ME  (2000)  Behavioural correlates of predation by tiger (Panthera tigris), leopard (Panthera pardus) and dhole (Cuon alpinus) in Nagarahole, India.  Journal of Zoology 250: 255-265

Khan I  (1936)  Association between a leopard and a tigress.  Journal of the Bombay Natural History Society 39: 155-156

Nagata J, Aramilev VV, Belozor A, Sugimoto T, McCullough DR (2005)  Fecal genetic analysis using PCR-RFLP of cytochrome b to identify sympatric carnivores, the tiger Panthera tigris and the leopard Panthera pardus, in far eastern Russia.  Conservation Genetics 6: 863-866

Palomares F, Caro TM  (1999)  Interspecific killing among mammalian carnivores.  American Naturalist 153: 492-508

Seidensticker J  (1976)  On the ecological separation between tigers and leopards.  Biotropica 8: 225-234

Schaller GB  (1967)  The deer and the tiger.  University of Chicago Press, USA

Sankar K, Johnsingh AJT  (2002)  Food habits of tiger (Panthera tigris) and leopard (Panthera pardus) in Sariska Tiger Reserve, Rajasthan, India, as shown by scat analysis.  Mammalia 66: 285-289

Støen OG, Wegge P  (1996)  Prey selection and prey removal by tiger (Panthera tigris) during the dry season in lowland Nepal.  Mammalia 60: 363-373

Sunquist F, Sunquist M  (2002)  Tiger moon: tracking the great cats of Nepal.  University of Chicago Press, USA

Venkataraman AB, Arumugam R, Sukumar R  (1995)  The foraging ecology of the dhole (Cuon alpinus) in Mudumalai Sanctuary, southern India.  Journal of Zoology 237: 543-561

Information about this Review

The author is:  Dr. Paul D. Haemig (PhD in Animal Ecology)

The photograph of the tiger was taken by Lee Anne Kortus of Porter, Texas.

Other contributions:  This paper was improved using information submitted by a tiger expert from Vietnam who chooses to remain anonymous.

The proper citation is:

Haemig PD  2007   Sympatric Tiger and Leopard.  ECOLOGY.INFO #22

If you are aware of any important scientific publications about sympatric tiger and leopard that were omitted from this review, or have other suggestions for improving it, please contact the author at his e-mail address: 

paul.haemig {at} hik.se

© Copyright 2003-2007 Ecology Online Sweden.  All rights reserved.

reviews
poems
anthology

advertise
about
students
teachers
instructions
  for authors

donate photos
  and artwork

translate

copyright
disclaimer
faqs
write us

home

bahasa melayu
castellano
deutsch
filipino
français
português

 

For

 those

 who want

 up-to-date

 info

 NOW!

ecology.info