Updated: 6 August 2021
Throughout much of its range in Asia, the tiger (Panthera tigris) is sympatric with the leopard (Panthera pardus). Because both cats are large 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 tiger and leopard when sympatric (i.e. live in the same area) and summarize the important differences between them that reduce competition and allow coexistence.
To fully understand coexistence, however, we must also consider 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 also eats some of the same ungulates. Although smaller in size than the two big cats, the dhole hunts in groups and so is an especially formidable predator.
Because the tiger, leopard and dhole often live in the same areas and eat many of the same prey, we will review studies not only of the tiger and leopard but, when available, of the dhole as well, to understand how all three predators differentiate themselves to avoid violent encounters and competition for food.
Differences in Body Size and Prey Size
The tiger averages four times heavier in weight than the leopard (Seidensticker 1976). The leopard averages three times heavier than the dhole (Sunquist & Sunquist 2009, Sillero-Zubiri 2009).
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 usually selected prey weighing 31-175 kilograms.
At the Anamalai Tiger Reserve, southern India, the average size of prey caught by tiger was 92 kilograms, by leopard 37 kilograms, and by dhole 36 kilograms (Kumaraguru et al. 2011).
Predation on Elephants and Rhinos
Adult elephants and rhinoceroses are so large that 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 young Indian Rhinoceros (Rhinoceros unicornis) whose mother was killed by a tiger.
Tigers prey more frequently upon elephant calves and rhino calves than do leopards. For example, 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 wild ungulates (hooved animals)
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 Eurasian wild pig (Sus scrofa). All three carnivores preyed on each ungulate species, but there were differences in frequency. Tiger preyed more frequently on 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 generally immune to tiger predation (Johnsingh 1983).” Also, sambar spend more time in 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 pigs, and dhole rarely killed them, but tigers readily killed large adult male wild pigs, which generally live alone (Johnsingh 1983). Similar results were found at Bardia National Park in Nepal. There, the most common ungulates (Chital and Hog deer Axis porcinus) were taken less frequently, and the rarer wild pig more frequently, than predicted by their abundances (Wegge et al. 2009).
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) hypothesized that the greater predation on male ungulates could be explained 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 preyed upon male sambar more than female sambar (Johnsingh 1983). Dhole also preyed more upon 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 Bengal gray langur (Semnopithecus entellus) is a leaf-eating monkey. In the tropical forests of Nagarahole National Park, remains of this langur were found in 38% of leopard scats, 19% of tiger scats and only 1% of dhole scats (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. Remains of another monkey, the rhesus macaque (Macaca mulatta), were found in 13% of leopard scats, but in only 4% of tiger scats (Bhattarai & Kindlmann 2012). Similar results were found in the Suklaphanta Wildlife Reserve, also in Nepal (Lovari et al. 2015).
The more frequent predation by leopard on monkeys 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 monkeys can detect it easier than tiger or leopard, which silently 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. Among the many hypotheses that need to be considered is the possibility that some of the leopard scats were misidentified as tiger scats.
Predation on Smaller Prey
At the Sariska Tiger Reserve, Sankar & Johnsingh (2002) found rodents (mainly Indian gerbile Tatera indica) and insectivores (mainly grey musk shrew Suncus murinus) 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 Indian hare (Lepus nigricollis) and Indian crested porcupine (Hystrix indica) than did tiger (Karanth & Sunquist 1995).
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.”
Dhole leave most of their kills in the open, while both the tiger and leopard usually hide their kills in dense cover (Karanth & Sunquist 2000). An exception occurs with gaur carcasses, which are too heavy to be dragged, and consequently are often left in the open by tiger (Karanth & Sunquist 2000).
Although the tiger, leopard and dhole hunt at all times of day or night, 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 three 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 a move-and-stop hunting strategy, 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).
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, tiger usually attacked in more open settings, 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 kills less 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).
The leopard used the burned areas more frequently than the 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 there 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, however, 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 reasons for 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.
For example, 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 two 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 switched to hunting the medium-sized prey eaten by leopard. The resulting diet overlap led to increased encounter rates with leopards and increased levels of aggression toward them.
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 tiger, leopard and dhole.
In contrast, large ungulates are scarce in Bardia, causing tiger to shift to the smaller ungulates preferred by leopard (Støen & Wegge 1996; Karanth & Sunquist 2000).
If the scarcity of large ungulates in Bardia is the result of illegal poaching by humans, then it could be legitimately argued that such criminals indirectly cause the leopard depredations on neighboring farms and villages, since they force tigers to shift to smaller prey, taking over the food resources of leopards and displacing the latter cats to seek food in nearby farms and villages (see also Shehzad et al. 2015).
In Manus National Park (India), Lahkar et al. (2020) studied sympatric tiger, leopard and dhole in environments where prey was abundant, but human influences altered the spatial and temporal behavior of the prey. They found that all three predators segregated through fine-scale spatio-temporal avoidance rather than by displaying population-level changes in space-use, activity patterns or food habits.
Interactions in China and Russia
So far in this review, we have looked at sympatric tiger, leopard and dhole on the Indian subcontinent. However, all three species also occur in China and southern Russia. Here, at the northernmost limits of their distribution, field research gives us further insights into their habits and differences.
Coexistence between these carnivores in the North is determined not only by dietary differences, but also by differences in habitat use and differences in the time of day they are active. However, the three predator species also appear to respond differently to human disturbance, and these different responses can result in displacements that alter niche differences (Yang et al. (2018b; Manlick & Pauli 2020; Seveque et al. 2020).
In NE China and SE Russia (The Amur region), both the tiger and the leopard feed on many kinds of animals, large and small (Sugimoto et al. 2016; Yang et al. 2018). However most of their prey are three species of wild ungulates: sika deer (Cervus nippon), Eastern roe deer (Capreolus pygargus) and Eurasian wild pigs. Like the tigers and leopards of the Indian subcontinent, tigers here prey upon Eurasian wild pigs more frequently than do leopards (Sugimoto et al. 2016; Yang et al. 2018a).
In addition, Amur tigers are recorded more frequently in valleys (Carroll & Miquelle 2006), while Amur leopards are recorded more frequently on ridges (Yang et al. 2018b). Amur leopards are also more active during daytime than Amur tigers, the latter appear to avoid humans when the latter are most active. (Yang et al. 2018b, Li et al. 2019).
In the Land-of-the-Leopard National Park (Russia), Matiukhina (2020) found that when human impacts on the park were minimized, there was adequate food and space for both Amur tigers and Amur leopards to coexist. She found no evidence that tigers displaced leopards or vice-versa. On the contrary, when so protected from human impacts, both species thrived, and “the number of observed leopard litters of all ages increased positively with local tiger density (Matiukhina 2020).”
This remarkable discovery suggests that endangered Amur tigers might actually enhance the survival of endangered Amur leopards. For example (and this is speculation by the author of the present review), if fearful wildlife fleeing from tigers move unsuspectedly toward leopards, they might be caught more frequently by leopards. Another possibility is that tiger presence protects leopards in some way.
While this discovery is hopeful news, Matiukhina (2020) warns that the current positive relationship of the two big cats in the Land-of-the-Leopard National Park could change to negative if tigers are prevented from dispersing from the park to surrounding areas, causing tiger overpopulation that could harm the leopards. She points out that if both the leopard and tiger are to recover, liveable areas outside the park are needed where surplus individuals of both cat species can disperse to and make new homes.
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Information about this Review
The author is: Dr. Paul D. Haemig (PhD in Animal Ecology)
Other contributors: This paper was improved using information submitted by a tiger expert from Vietnam who chooses to remain anonymous.
The proper citation is:
Haemig PD 2021 Sympatric Tiger and Leopard ECOLOGY.INFO 22
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