Haemig PD  (2012)  Sympatric Tiger, Leopard and Dhole.  ECOLOGY.INFO 22

Sympatric Tiger, Leopard
and Dhole

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 reduce competition and 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 some of 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 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 & 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."

At the Anamalai Tiger Reserve, southern India, the average size of prey caught by tiger was 92 kilograms, for leopard 37 kilograms, for dhole 36 kilograms (Kumaraguru et al. 2011). 

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.  A few years ago, it was reported that the World Wildlife Fund in Nepal was 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 than do leopards.  In the tropical forests of Nagarahole National Park, Karanth & 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 Ungulates

In the tropical forests of Nagarahole National Park, Karanth & Sunquist (1995) found that five species of ungulates comprised 89-98% of the diet of sympatric tiger, leopard and dhole.  The five ungulates were gaur (Bos gaurus), sambar (Rusa unicolor), chital (Axis axis), barking deer (Muntiacus muntjak) and wild pig (Sus scrofa).  All three carnivores preyed on each ungulate species, but there were differences in frequency.  Tiger preferred gaur and avoided barking deer, 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).  Leopards did not kill wild pig, and dhole rarely killed it, but tigers readily killed large adult boars, which live alone (Johnsingh 1983).

A later study at the Bandipur Tiger Reserve compared biomass of prey consumed by the three predators (Andheria et al. 2007).  The tiger's diet consisted of 42% gaur, 31% sambar, 19% chital, 4% wild pig.  The leopard's diet consisted of 56% chital, 15% gaur, 9% wild pig, 8% sambar.  The dhole's diet consisted of 78% chital, 13% sambar, 4% wild pig, 1% gaur.

At the Anamalai Tiger Reserve, southern India, tigers preferred to prey upon sambar, Nilgiri tahr (Hemitragus hylocrius) and gaur, avoiding chital and wild pig (Kumaraguru et al. 2011.  Leopards preyed most frequently on black-naped hare (Lepus nigricollis) and mouse deer (Tragalus meminna), followed by sambar and nilgiri tahr, and avoided wild pig, chital and gaur.  Dholes preyed most frequently on barking deer, sambar and mouse deer, avoiding wild pig, chital, gaur, mouse deer and nilgiri tahr

Predation on different sexes and age groups

At Nagarahole, Karanth & 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.

Karanth & Sunquist (1995) 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 & 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 & Sunquist 1995).   Furthurmore, langur was under-represented in the diets of both the tiger and dhole, indicating avoidance (Karanth & Sunquist 1995).

Similar results were found at Bandipur Tiger Reserve.  Langurs made up 7% of the prey biomass consumed by leopards, but less than 1% of the prey biomasses consumed by tiger and dhole (Andheria et al. 2007). 

These results are further corroborated by findings from a study in Bhutan.  There, langur remains were found in 8% of leopard scats but not in any tiger or dhole scats (Wang & Macdonald 2009).

At Chitwan National Park in Nepal, Langur remains were found in 17% of leopard scats, but in only 7% of tiger scats.  Rhesus monkey remains were found in 13% of leopard scats, but in only 4% of tiger scats (Bhattarai & Kindlmann 2012).

The more frequent predation by leopard on langur can be explained by the fact that the leopard climbs trees better than the tiger (Karanth & 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 & 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 & Johnsingh 2002).  The reason for these different results is unknown and worthy of further investigation.

Predation on Smaller Prey

At the Sariska Tiger Reserve, Sankar & 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% of 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 & Sunquist 1995).

Food Caching

The leopard differs from the tiger and dhole in that it often takes the carcasses of animals it kills up into the trees to eat and cache for future consumption.  There 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 & Sunquist 2000).  Gaur carcasses, however, which are too heavy to be dragged, are often left in the open by tiger (Karanth & Sunquist 2000).

Activity Periods

Although the tiger, leopard and dhole hunt round the clock, the dhole differs from the two cats by being mainly diurnal (Johnsingh 1983; Venkataraman et al. 1995; Karanth & 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 & Sunquist 2000).  The dhole was the only one of the 3 predators that killed prey frequently in the afternoon (Karanth &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 & Sunquist 2002, p. 107)."

In a tropical forest on the Malay Peninsula, the tiger was found to be mainly nocturnal, while the leopard was mainly diurnal (Azlan & Sharma 2006).

Microhabitat Use

At Nagarahole, tiger attacked 81% of their prey in "dense or moderate cover (Karanth & 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 & 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 & 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 & 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 & 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 & Sunquist 2002).

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 & Sunquist 2002).

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 & Sunquist 2002).

Utilization of Roads

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

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 & 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 & Sunquist 2000).  Leopard predation on dhole is reported from both Bandipur (Johnsingh 1983, 1992) and Nagarahole (Karanth & Sunquist 1995).

At Bardia National Park, Nepal, Støen & 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.

At the same park, Odden et al. (2010) reported that when the large ungulate prey that tigers need became scarce, tigers were driven into competition with leopards for medium-sized prey.  These researchers concluded, "Low abundance of large ungulate prey decreases foraging efficiency of tigers, leading to increased energetic stress and aggression towards leopards; and increased diet overlap due to scarcity of large prey leads to increased encounter rates and increased levels of interference competition."

Similar findings are reported for Kanha National Park, central India (Schaller 1967) and Rajaji National Park, northern India (Harihar et al. 2011).  At Rajaji, a population of tigers recovering from near extinction grew and took over the optimal habitats of the park.  Leopard density in the areas taken over by tigers declined to less than one-fourth its previous level, and many leopards shifted to tiger-free areas outside the park where they then fed on domestic livestock.

Killing and displacement 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 Bardia National Park does not occur (Karanth & Sunquist 2000).  Nagarhole has an abundance of both large and medium-sized ungulates, providing adequate food for both tiger and leopard.  In contrast, large ungulates are scarce at Bardia, causing tiger to shift to the smaller ungulates preferred by leopard (Støen & Wegge 1996; Karanth & Sunquist 2000).

Sunquist & Sunquist (2002) 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."

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