THE AFRICAN LEOPARD

The leopard, more than the lion and the cheetah, is the quintessential big cat because of its lifestyle and behaviour which is so typical of felids. Very adaptable, it was once widespread throughout Africa from the Maghreb to South Africa with the exception of the Sahara. Today, it has almost disappeared from the Maghreb and has become extremely rare in West Africa. It remains relatively common in eastern and southern Africa but is threatened in central Africa due to the disappearance of its habitat and its prey due to the bushmeat trade. A victim of its beauty, the leopard is a prime target for illegal wildlife trafficking to Asia where its bones and fur are highly prized by wealthy customers. The African leopard is itself a subspecies of the leopard, but its appearance varies according to region. Leopards in Somalia and the Cape region, for example, are much smaller than average, weighing around 30 kg, while leopards in southern Africa are more massive than in the rest of Africa. Specimens from East and Central Africa are also among the largest on the continent. Unlike some Asian leopard subspecies that may be melanistic, notably the leopards of Malaysia, Indochina, Java and more rarely India and Sri Lanka, melanistic leopards (also called black panthers) are very rare in Africa.

The leopard can be distinguished from the cheetah, with which it is often confused, by a more massive and muscular body, shorter and more muscular legs, a massive head and neck without black teardrop-shaped spots under the eyes, and a coat studded with rosettes, unlike the cheetah's coat, which is studded with solid black spots. Their behaviour and habits are also opposite. The leopard hunts by ambushing its prey, which it subdues by brute force, whereas the cheetah uses its speed. The cheetah is also more sociable than the leopard, which leads a solitary life. The leopard is also better equipped than the cheetah to deal with interspecific competition.

Female leopard (Sabi Sand, South Africa)

MEASUREMENTS

Length (without the tail): F= 95-127 cm ; M= 92-123 cm
Shoulder height: 60-70 cm
Weight: F= 20-50 kg (up to 60 kg for biggest) ; M= 30-80 kg (up to 80-90 kg for the biggest)

ECOLOGY

Habitat: The leopard is the most adaptable of the big cats and has colonised all areas of the African continent from semi-deserts like the Kalahari to the rainforests of the Congo Basin and even the outskirts of large cities like Nairobi. It is only absent from true deserts like the Sahara. It is the only large African predator found in the rainforest of the Congo Basin where it is the apex predator. The leopard is also present in all savannah ecosystems, but with a preference for woodland savannahs, miombo, and mosaics of woods and clearings. It is scarcer and vagrant in short grass savannahs and grasslands. In this type of environment, it often uses rocky outcrops such as the kopjes of the Serengeti as a refuge. It can also be found in mountainous areas such as the Kilimanjaro, the Bale Mountains in Ethiopia up to 4000 m in altitude.

Food: The leopard is able to colonise so many environments because it is able to feed on a wide range of prey species. Its diet is the most diverse of any cat, including the lion. 92 species have been recorded as prey in sub-Saharan Africa alone.  Indeed, its average size allows it to survive by feeding on small preys such as rodents, fish, birds and small antelopes that would not sustain a lion and which often figure in the leopard's diet. On the other hand, it is still powerful enough to take down large ungulates weighing three times its weight if the need arises. One famous example is the male leopard that managed to kill a 900 kg adult Cape eland after throwing himself on its back from a tree. There are also documented cases of leopards successfully attacking adult zebras, greater kudu (including bulls), waterbucks, hartebeests, and topcis (and affiliated). However, this remains unusual and in general the leopard prefers to attack smaller prey weighing 20-80 kg in average with a preference for small and medium-sized ungulates and primates. Preferred prey includes impala, bushbucks, Grant's gazelle (and affiliated), Thompson's gazelles, reduncas, nyala, puku, kob; lechwe, sitatunga, lesser and greater kudu (especially females and juveniles), various species of duiker, warthogs and bushpigs, zebra foals and sub-adults, as well as juveniles and sub-adults of large antelopes comprising sables and roans, gemsboks, wildebeests, topis, hartebeests and waterbucks. More rarely, the leopard may attack adults of these same species. In southern Africa, it seems that large female kudu, which can weigh between 120 and 200 kg, are more common prey than one might think, based on many existing photographs. There is also a video on YouTube showing a lame male leopard capturing effortlessly an adult zebra mare who was also injured after a crocodile attack.  One might ask why leopards do not attack large adult ungulates more often, as do the Himalayan snow leopard or the puma in the Americas, which are of similar size. It would seem that there is little point in a leopard attacking prey weighing more than 100 kg, as a 60 kg impala is sufficient to satisfy its food requirements for several days. On the other hand, attacking large ungulates such as wildebeest, waterbuck, gemsbok or sable is not without risk of injury and is not worth taking if dietary needs can be met with smaller, less dangerous and more manageable prey. It will be also much more difficult for a leopard to protect large prey from other predators. Indeed, a leopard can put its prey out of reach of other predators by storing it on a tree, its exceptional power allowing it to climb with a 125 kg prey in its mouth. Beyond this weight, the prey will have to remain on the ground and therefore remain vulnerable. If a lion or a group of hyenas or wild dogs come onto the scene,  the leopard will be obliged to abandon its prey, and the expenditure of energy in addition to the risks incurred to subdue a large prey will have been in vain. Of course, snow leopards and pumas also face kleptoparasitism from brown bears, wolves and perhaps even faras for the snow leopards and wolverines for the puma, but it seems that the density of these predators is less than lions and hyenas in African conservation areas or tigers and dholes in Indian conservation areas.  It seems therefore that in the presence of other predators, it is wiser for the leopard to specialise in small and medium-sized prey that are of less interest to the more  social predators such as lions, hyenas and wild dogs who need larger food intakes from bigger prey.

However, when other large predators are absent, it appears that leopards, like cheetahs, are more likely to hunt larger prey. In Sri Lanka, for example, leopards are more likely to attack adult sambar deer, which are the largest deer in Asia, equivalent in size to a greater kudu, whereas in India they prefer to prey on juveniles and sub-adults, leaving adult sambars to the tiger and dhole packs.  Similarly, in the tropical forests of Central Africa, where it is at the top of the food chain, the leopard does not hesitate to set its sights on adult gorillas, okapi, bongo and giant forest hog even though medium-sized prey such as sitatungas, duikers and primates remain preferred prey. Throughout Africa, the leopard is the main predator of all primates. Only the largest species such as gorillas (including silverbacks), chimpanzees, mandrills and various species of baboons are armed enough to resist a leopard. Baboons live in packs of up to several dozen individuals and are protected by adult males that can weigh up to 30 kg and  armed with canines as long as those of a leopard. Adult male baboons, sometimes joined by females, will stand together in the face of a leopard attack and in most cases manage to repel it. Chimpanzees and gorillas do the same, except that they are even more powerful and intelligent than baboons. Consequently, it is rare for the leopard to attack these primates head-on, as they may kill it in retaliation. The leopard usually waits until nightfall before attacking a lone individual that is lagging behind.

The leopard also attacks other predators such as pythons, medium-sized crocodiles, adult cheetahs and other smaller felids such as African golden cats, servals and caracals, lagging adult wild dogs, jackals and young hyenas. Other unusual prey includes aardvarks, adult ostriches and the calves of buffalo, giraffe and rhino, which it even manages to hoist into trees.

The leopard is a stalking hunter hich ambushes its prey. For medium-sized prey, such as ungulates weighing between 20 and 80 kg, once spotted, it approaches as close as possible in silence by crawling on the floor using the vegetation and any other feature enabling it to hide from its prey. Once it is about 5 metres from its prey, it accelerates rapidly to grab it by the rump or shoulders with its powerful front legs to bring it to the ground. It then grabs the prey by the throat to choke it with a fatal bite. For prey weighing more than 100kg, such as adult greater kudu, the leopard jumps on their backs - like lions when they attack a buffalo - and claws at them until they collapse under his weight, allowing him to make the fatal bite. The male leopard that killed the 900 kg adult eland proceeded this way and hanged on the eland until it collapsed. Sometimes the leopard grabs onto the necks as best he can until he finds an opportunity to make the killing bite. There are several photos of leopards doing this to bring down adult topis and waterbuckse. It seems that leaping from the top of trees onto its prey makes it easier to kill large ungulates, a Youtube video shows a leopard taking down a sub-adult zebra in this way in the Serengeti. It will also succeeded in hoisting it onto the same tree. When hunting prey weighing less than 20 kg, a simple bite on the neck is enough to finish off the prey. Once killed, the leopard drags its prey along the ground and takes it into cover so that it can devour it in peace without dusturbance. In areas with high concentrations of hyenas and lions that could steal its prey, the leopard hoists its prey up a tree out of reach of other predators, especially hyenas. This requires colossal strength. The leopard is able to hoist prey weighing up to 130 kg onto the forks of trees. The body of a young giraffe weighing 125 kg was found on a tree fork hoisted by a leopard. One YouTube video shows a leopard hoisting a young rhino up a tree, another shows him hoisting a sub-adult zebra. Hoisting a 60 kg impala is therefore not a problem at all. This ability to use trees to store prey and take refuge explains why the leopard is able to cope better than wild dogs and cheetahs with competition from other large predators.

Behaviour and social structure: The leopard is a solitary animal, which does not necessarily mean that it is asocial similar to the tiger. The male and female leopard only meet for mating, during which time they may stay together for several days. Like the tiger, the male leopard tolerates his own offspring and brief family reunions including the cubs and both parents may occur. Female leopards often tolerate the presence of their daughters from a previous litter, who sometimes establish their territory close to their mother's or even partially overlaps with it.  Apart from mating and female leopards with their cubs, leopards avoid each other. Males and females defend territories against other leopards of the same sex. The territories of males include those of several females. Adult leopards patrol their territories, marking them with faeces and by spraying trees and other prominent spots with urine and then scratching the ground with their hind legs. The territory is also marked by rubbing the head against trees. Like the lion, the leopard roars to indicate that the territory is occupied and to keep potential rivals at bay. The leopard's roar is very different from that of the lion and is similar to the sound of a saw on a tree. It can carry up to 3 km, thus avoiding confrontation. In the event of an encounter, a show of force involving a lot of growling, intimidating charges, hissing, snarling and exposure of the canine teeth is enough to get the weaker leopard to back down into its territory. If these displays of force are not enough, leopard fights can be very violent and result in injury or death to one of the combatants.

Leopards are rather active at night which they devote to hunting and patrolling their territory. They usually spend the day resting in the vegetation or on a tree.

Breeding behaviour: Capable of breeding all year round. The female usually initiates the courtship where she solicits the male on he tracks by rubbing her cheeks against his before moving into position for mating.  During this period they may hunt together and share prey. As with the lion, the leopard lightly grasps the female's neck during mating and jumps back at the end of the process to avoid the female's slap. The female gives birth to a litter of 1 to 3 cubs after a gestation period of 90 days. She gives birth in a cave hidden by vegetation. The young are born blind. They open their eyes after about ten days and come out of their hiding place after 6 weeks. They are weaned after three months. The mother protects them from predators and foreign male leopards that might kill them in order to mate with the mother and thus pass on their genetic heritage. As with the lion, infanticide is practised in leopards. However, she is obliged to leave them alone to go hunting, during which time they are very vulnerable. After a month, they are able to take refuge in the trees and thus protect themselves from predators when their mother is away. She calls them when she returns from the hunt and leads them to the killed prey. The young are independent after a year or a year and a half. At this time, the mother sometimes makes it clear to them aggressively that they must leave. Some young try to stay with their mother and still feed on her prey. Young females usually establish a territory close to their mother's, while young males roam further away. As with the young lions, this is a difficult time, especially as they are on their own. The young male will have to establish his own territory, which may mean that he has to oust an older male to steal all or part of his territory.

Predators and competitors: Lions are the only African land predators individually capable of killing a healthy adult leopard and therefore pose the greatest threat. Leopards often take refuge in trees when approached by lions that are not able to climb as high as a leopard. If cornered, the leopard will defend itself valiantly and get on its back to protect its vulnerable spine and deliver numerous slaps to the face of the attackers. Several Youtube videos show leopards successfully defending themselves even when surrounded by a whole pride of lions, even managing to injure a few of them and finally escaping by taking advantage of a moment of inattention. In one of those videos, a male leopard was feasting on the carcass of a greater kudu killed by lionesses in the Great Kruger before being surprised and surrounded by four or five of them. The lionesses tried to attack him, but the leopard got on his back and pawed at his assailants, injuring one on the snout. The lionesses only attacked the leopard when its attention was diverted by another and it was very clear that they were reluctant to attack it head on. The appearance of a hyena attracted by the noise allowed the leopard to escape. These videos show that lions are cautious or hesitant when attacking a leopard. This is even more true when the lion, male or female, is alone. The explosiveness, aggressiveness and agility of the leopard often allow it to keep its much larger opponent confused by the leopard's aggression allowing the leopard to flee before the lion catches up. In another video, probably shot in South Luangwa, Zambia, we see a male lion in its prime managing to approach a sleeping leopard which at the last moment wakes up in an explosive jump scare which surprised the male lion. The leopard, taking advantage of the latter's confusion, managed to escape. Two other similar videos show a leopard surprised by a lioness and another by a lion where the leopard's aggressiveness and explosive reaction allowed it to escape before the lion or lioness could react. The leopard is not always so lucky and sometimes falls to the lion's fangs when surprised. The lion does not eat the carcass of a leopard it has killed. It is more a matter of removing a competitor.

The relationship between leopards and spotted hyenas is even more complex as they are predators of more or less equal size and power depending on the specimen. Spotted hyenas are more competitors than predators. It is highly unlikely that a spotted hyena would want to attack a healthy adult leopard, assuming it is capable of doing so, unless the leopard is weakened or injured. However, spotted hyenas have been known to kill sub-adult leopards and cubs. A group of spotted hyenas would also dispatch a healthy adult leopard without difficulty and leopards take refuge in trees when hyenas are out in numbers. When there is no food involved and a leopard encounters a lone hyena or a small group, the two predators simply observe each other and keep a respectful distance from each other. This suggests that the adult leopard does not consider the hyena as an immediate threat to its life, unlike the lion.

Things become much more complicated when a carcass is involved. In this case hyenas are much more enterprising and even a lone hyena will usually manage to steal its prey from a leopard. The leopard sometimes tries to intimidate the hyena by growling and clawing at her, but the hyena is able to take it and call on the leopard's bluff. Observers are sometimes surprised that the leopard offers so little resistance. This can be partly explained by the fact that the hyena has the most powerful jaw of any land carnivore. Although the hyena's jaw is not designed to deliver a fatal bite, the leopard is likely to be injured in a fight with a hyena, especially if there are several of them, which would render it unable to hunt and condemn it to a slow death. It is therefore wiser to leave the prey to the hyena and hunt another prey rather than risk an incapacitating injury. For the same reasons, the leopard usually gives in to striped hyenas and solitary brown hyenas, even though they are smaller than the leopard. However, there is a video on Youtube of a leopard violently attacking a brown hyena that is trying to steal its prey by biting it in the throat.

Yet, male leopards, and more rarely females, are sometimes able to resist hyenas' attempts to steal their food and even manage to scare them away. Theodore Bailey even reports a case where a male leopard in the Kruger killed two adult spotted hyenas that were trying to take over the warthog he had just killed. In a report from Mala Mala in South Africa, a male leopard and a young female leopard are seen fending off lone hyenas that were trying to steal their food. Also in Mala Mala, a leopard almost killed an adult spotted hyena by grabbing it by the throat while the hyena was eating an impala. The outcome of confrontations between leopards and hyenas depends on the size and sex of the parties involved, the number of hyenas present and the circumstances. Female spotted hyenas are larger than male hyenas and are more powerful. The average spotted hyena is also larger than the average male leopard, but larger leopard specimens are larger than the largest spotted hyena specimens. Spotted hyenas alone in groups are therefore usually able to steal their prey from leopards, but this is by no means an unalterable rule. Male leopards, which are larger and more aggressive than female leopards, can sometimes repel one or two spotted hyenas. Female leopards will also attack hyenas head-on if they are attacking their cubs and will manage to drive them away. Both species will also attack young left unprotected.

The relationship with wild dogs is also complex. A leopard is much more powerful than a single adult wild dog, and a leopard will sometimes kill a wild dog lagging behind. But it is relatively rare for a wild dog to be without its pack. A pack of wild dogs with more than three or four members is usually able to scare a leopard away by chasing it into a tree. A pack of wild dogs, like packs of dholes in Asia, can kill a leopard by the weight of their number. This is particularly the case when the leopard tries to attack the cubs or a straggling adult, or tries to steal the wild dogs' prey. Wild dogs deprive leopards of their prey more than the reverse. Leopards may sometimes be able to steal their prey from groups of two to three dogs but generally thew generally avoid packs of wild dogs as well as groups of hyenas by taking refuge in the trees but the lone or straggling wild dog must beware.

The leopard almost invariably dominates cheetahs, especially solitary ones and even coalitions. Cheetahs are considered as prey that the leopard hoists into the trees as if it were an antelope. A female cheetah will however not hesitate to confront a leopard to protect her cubs by diverting its attention and using her speed to escape when the leopard turns against her. There are also two cases where two male cheetahs managed to scare away a leopard and another case where five brother cheetahs harassed a male leopard  and treed him but these are exceptions.

The leopard also dominates and kills all predators smaller than itself which includes all small and medium sized felids such as servals, caracals and golden cats, jackals, aardwolves, mongooses, civets but also African rock pythons and small and medium sized crocodiles.

Large Nile crocodiles (over 4 m) are another matter. Alongside lions, they are the only predators in Africa capable of subduing singlehandedly a healthy adult leopard. Theodore Bailey reports that an adult male leopard, the famous hyena killer, was killed by a Nile crocodile.

Some prey such as baboons and chimpanzees are also capable of killing an adult leopard using their overwhelming numbers, although this is rare. The leopard usually retreats once spotted and the primates mock charging and shout but avoid direct contact with the predator. Leopards can also injure themselves, sometimes fatally, when trying to tackle dangerous prey such as warthogs, porcupines and large antelopes such as waterbucks, gemsboks, sables or zebras, which explains why these large animals are rarely on the menu.

Best places to see them: Unpredictable. Although present in all the major protected areas of eastern and southern Africa, the leopard's nocturnal habits, elusiveness, not to mention its ability to hide in the vegetation, make it very difficult to observe. In some reserves where they are used to the presence of vehicles, it can be relatively easy to observe them. The private reserves adjacent to the Kruger probably offer the best chances and it is possible to see several leopards in one day which is rarely possible elsewhere in Africa. After the private reserves in South Africa, the South Luangwa Park in Zambia has an excellent reputation where leopard viewing is common in perhaps more authentic surroundings than in South Africa. The Seronera region of the Serengeti and the Masai Mara are also great places for leopard watching. Three days on safari usually guarantees at least one sighting. Moremi also offers opportunities although I have never seen a leopard in Botswana nor in Zimbabwe. As is often the case with wildlife viewing, it's all about luck and being there at the right time.