© Lions at dusk © Heinrich van den Berg
One of Africa's most unforgettable experiences is hearing the roar of a lion at night. When Ingonyama (Swazi) or Nghala (Shangaan) announces his presence with a deep roar that reverberates through the dark bush, everything within a five-kilometre radius pauses to take note.
A thousand years ago, they roamed as far afield as southern Europe, the Middle East and Asia, but they are now found mainly in Africa and are generally restricted to the bigger game reserves.
Lions have always had an association with royalty and leadership. Their power is reflected in their impressive size and the fact that their lifestyles allow them to sleep a lot - up to 18 hours a day. They are awesome animals, with males weighing up to 225kg and females up to 150kg. The lion's average lifespan in the bush is probably around 15 years.
Lions are the biggest, and most social, of the African cats, living communally in prides. Pride leadership often shifts between several individual animals - male and female - but the social structure of the pride hinges around the bond between related lionesses, who collaborate in all tasks, from raising cubs to hunting.
Male lions come and go - often in spectacular battles over territory or individual dominance - but the pride cohesion remains unaffected, firmly under female control.
Lions favour open woodlands and thick scrub, the type of landscape that allows them to get as close to their prey as possible without being seen. In Kruger, the best chance of lion sightings is where the big game herds are. As a rule, they hunt mostly at night and rest during the day but are often active at dawn and dusk and on cooler days. During the day they rest in thorn thickets, often near water holes.
Lions are believed to feed every three or four days, and need on average between 5kg and 7kg of meat a day. But they can go without food for more than a week and then tear into prey, eating up to 50kg of meat at a time - that's almost a quarter of the animal's body weight.
Lions hunt either collaboratively or by themselves. Collaborative hunting usually involves the males approaching the intended prey upwind with the intention of driving it towards lionesses hiding in the bush downwind. Lions are not as fast as most of their prey so they rely heavily on the element of surprise when hunting.
They will attempt to get to within 30m of their prey before charging. They don't have the inclination for a long chase and will not pursue their prey very far if the first attack fails.
In a successful hunt, the prey is knocked off balance, dragged down and then killed with a bite to the back of the neck or the throat. In some cases, a kill can be a bloody, drawn out procedure. Buffalo have been known to fend off lion attacks for hours before succumbing to loss of blood and energy. The strongest male lion will eat first, followed by other members of the pride. Lionesses will feed themselves first, with cubs getting the scraps.
Lions sometimes become the victims of their intended prey. There have been instances in which lions have been killed by giraffe, buffalo, kudu, snakes and even porcupines.
What do Lions Eat?
The short answer is quite a lot. In Kruger, lions have a broad diet with 37 animal species on the menu, including ostriches, quelea nestlings, tortoises and small crocodiles. Their preference is for buffalo, giraffe, zebra and wildebeest among the bigger animals, and porcupine and warthog as smaller game.
Natural history artist Charles Astley Maberly - who rode through the Park on his bicycle sketching the wildlife in the 1920s - said that, in his experience, lions had a particular preference for waterbuck. This was confirmed by a 1960s study by former Kruger Chief Ranger Tol Pienaar who measured the kill rate of lions in the Park against the relative abundance of particular prey.
He found that, although lion killed relatively few giraffe, giraffe meat accounted for almost a third of the average lion's diet. His study also found that, contrary to popular belief, lions are not so fond of impala.
Kruger researchers suggest male and female lions may also have different prey preferences, with males being more disposed towards hunting buffalo, while lionesses prefer zebra or wildebeest.
Another fact not commonly appreciated is that lions are not just hunters, but scavengers as well, often chasing smaller predators - like cheetah - off their kills. In some instances, up to 50% of a lion's diet can come from scavenging rather than hunting live prey.
Lions on a Killing Spree
During periods of drought, lions sometimes go on a killing spree when they come across herds of weak animals. During the particularly bad drought of 1964, a pride of lions killed 15 buffalo near Punda Maria - far more than they could have eaten.
The territorial behaviour of lions is complicated because prides split up and re-unite, and hunting grounds shift as the seasons change and the game moves around. At any one time, a pride's territory in Kruger measures approximately 10 square kilometres. Territories may be defended vigorously by both males and females, but there are instances when prides share the same overlapping hunting zone yet deliberately avoid confrontation.
Clashes occur between prides when game migrations force lions to move beyond their territories in search of food, or when nomadic males challenge pride hierarchies. Fights can take the form of symbolic aggression displays and/or ferocious physical clashes that often result in the death of one or more participants.
If the challengers win, the defeated males are expelled and, in turn, become nomads themselves. The lionesses accept the new regime. Usually the conquering males kill all the cubs of the conquered pride. Within days of this infanticide the females come into oestrus and are ready to raise a new litter of the conquerer's cubs.
Scientists believe that territorial challenges are good for the survival of the species - they ensure diversity in the gene pool and dominance of the strongest genetic characteristics.
Lionesses typically give birth to litters of between two and four cubs. They are kept hidden in the bush for about six weeks. During this time, the lioness hunts exclusively for her cubs. Males provide no help. By the time they are two years old, cubs will have learnt to hunt for themselves.
At this time, male cubs are expelled from the pride while females are nurtured within it. These young males often form groupings of nomadic bachelors and either find new territories or challenge males in existing prides, and so the cyclical struggle for dominance goes on.
Anatomy of a lion kill
Lions hunt either alone or in prides. In collaborative hunts, it is usually the lioness who initiates the kill. Lions stalk their prey and, when close enough, attempt a short charge on their prey, trying either to pounce on their target or knock it over. They then kill their victim by breaking its neck or suffocating it by clamping their jaws around its throat.
The stomach is usually the easiest point of entry into the carcass, and this is the route most often taken by lions. It also gives them direct access to some of the most nutritious parts of the body, such as the kidneys and liver of the prey. Lions usually rest after an initial feed, lying a short way away from the carcass so that they can still defend their kill against scavengers.
In a short time vultures are certain to begin to arrive. The first are usually the white-backed vultures and then come the lappet-faced and others. Scavengers like hyaenas and jackals will be attracted by the vulture activity and will patiently wait at a safe distance until the lions have had their fill. It can take over 24 hours before lions abandon their carcass and spectacular fights may occur among scavengers for the last scraps of food.
Read more about Kruger's Big Predators