Field guide to lions

There is probably no other wild species as intertwined with human culture, history and evolution than the king of the beasts and there is good reason for that, says conservation group LionAid. 

No other animal is as well represented in symbolism, iconography, imagery, statuary, allegory, literature, totems, fables and art as the lion. Lions are symbols of nobility, bravery, courage, steadfastness and loyalty. They are represented in French cave paintings and as ivory carvings dating back 32,000 years, in San rock paintings in Namibia and Libyan petroglyphs dating back 6,000 years. The Egyptians loved lions – representing them as the massive Sphinx, the war goddess Sekhmet, and the Nubian gods Maahes and Dedun.

Narasimha, a lion avatar of Vishnu, is known as the Great Protector in Hindu culture, and lions form the emblem of India. Lions protected the gates of the Imperial Palace in Beijing and they even have a nation named after them – Singapore. Lions were considered the King of Beasts in Middle Age Bestiaries and hence feature on many coats of arms. Few cities today do not have lion statues, and their form is a favourite for company logos and names of sports teams.

Why are lions so interwoven with human culture? Doubtless because our distant ancestors held these formidable predators in great respect. As long ago as 3.5 million years, lions and hominids co-occurred in Tanzania. When Homo sapiens left Africa, they found lions already established in Europe and Asia. These animals had tremendous impact on the skills and intelligence of our species, and our protection from them was based on developing ever better weapons, secure housing and the use of fire. Wherever humans went, lions were there before them – Panthera leo spelea occurred in Eurasia and Panthera leo atrox ranged from Canada to Peru – both these lion subspecies only went extinct about 10,000 years ago. Cave paintings, ivory carvings and clay figurines imply early religious rituals involving lions – a respected predator and formidable competitor for the same species hunted for food by early man.

Despite such respect, lions have been eradicated from much of their previous geographic range. Once one of the most widely distributed terrestrial mammals, they are now only found in numbers in a few locations in Africa and one small remnant population in western India. The evolution of weaponry from primitive spears to bows and arrows and firearms, the use of domesticated animals such as dogs and horses to hunt lions, and the development of fast conveyances such as chariots and motor vehicles all contributed to the annihilation of lions wherever they posed a threat to humans and livestock. Eradication continues to this day, and has already seen many local extinctions presaging a catastrophic future for one of our most iconic yet vulnerable predators.

Lion subspecies

Based on morphology, distribution, mane colour and size in males and perhaps habitat adaptations, no fewer than eight subspecies of Panthera leo have been described in the past. These include Panthera leo persica, the Indian lion, Panthera leo leo, the Barbary lion that ranged from northern Africa into Europe (extinct in the 1920s, though a few might have survived wild in Morocco until the 1960s – captive descendants are in Moroccan and European zoos); and Panthera leo melanochaita described from South Africa. 

Recent genetic analyses indicate that western and central African lions are significantly different genetically from those in southern and eastern Africa and more closely related to the Indian lions. Therefore, three lion subspecies have been proposed – Panthera leo persica, Panthera leo leo in western/central Africa and formerly northern Africa, and Panthera leo melanochaita in southern and eastern Africa. 

Where do lions live?

Lions are found in scattered populations in sub-Saharan Africa. Because they are difficult to count accurately, population numbers are usually based on estimates. Today, approximations range from about 15,000 to 30,000 lions on the African continent, but only about 650 to 800 lions in all of western Africa. Some countries like the Congo Republic, Ivory Coast and Ghana have recently joined the ranks of 25 African nations where lions have gone extinct. In Senegal, Nigeria and Malawi at most a few score remain. Only five long-term viable populations remain in Africa, three in Tanzania/Kenya, one in South Africa, and one in Botswana/Zimbabwe. Current lion numbers are in great contrast to estimates of over 200,000 in the 1960s, meaning that in the past 50 years, lions in Africa have decreased by about 90 per cent.

Few people realize that there are fewer lions today than white rhinos, polar bears, chimpanzees, orangutans and a host of other endangered species. 

Lions’ physical characteristics

Lions are the second-largest wild cat species in the world (tigers have longer bodies but lions stand higher at the shoulder). Lion colouration varies among populations and individuals from light buff to a darker orange-brown. Underparts are generally lighter in colour. Lion cubs are born with dark brown rosettes (spots) on their bodies but these generally fade with age. Lions are the only cat to display sexual dimorphism in that males develop manes as they grow older. Such manes vary in size – some males might only have manes that cover the neck and chest, while others (especially lion males in captivity) can develop manes that extend along their backs and under their bellies. The mane colour varies from blond to dark brown to black.

Another unusual character among lions not seen in other cat species is the development of a tuft at the end of the tail.

Adult male lions vary in weight between 150-250kg, and females from 120-180kg. The heaviest recorded male lion came from the Colchester Zoo in the UK and weighed 375kg. While lions in captivity can live for more than 20 years, wild males rarely reach an age of 10-11 years and females 14 years.

What do lions eat?

Lions take a great diversity of prey, varying in size from small birds (Quelea) to catfish to adult giraffes and elephants. However, prey taken depends to a very large extent on habitat, seasonal movements of prey and species availability, as well as preferences among lion prides. Differences in diet can be very localised. In neighbouring prides in the Okavango Delta of Botswana, one hunted and killed many baboons, having learned that when baboons escape into trees they can be panicked to jump down again if a lion leaps up at the trunk. That same pride rarely hunted giraffes. The next-door pride never hunted baboons but had developed expertise at bringing down even large adult male giraffes.

As a generalisation, smaller prey such as warthogs, gazelles and impalas are hunted by solitary lions, while larger prey such as zebras, wildebeest, buffalos and giraffes require a group effort by many lions cooperating in a hunt.

Lion hunts are characterised (for some prey species) by long stalks, stealth, and then short but fast chases. Lions have little endurance and most attempts at catching nimble prey end in failure. 
Lions will hunt at any time of day, but concentrate their efforts around dusk and dawn and some parts of the night depending on the phase of the moon. Studies have shown that nights with a full moon mean poor success rates.

Lions also derive a significant part of their diet from scavenging and taking over kills made by other predators. They are extremely alert to descending vultures, at times following them for considerable distances to find a carcass. Likewise, lions are highly tuned to noises made by hyenas and wild dogs on their kills and will steal their prey. In Namibia lions patrol beaches to find washed-up carcasses of seals and other marine animals.

Lion behaviour

Lions are unusual among cats in that they are social. The social unit is the pride, a grouping involving stable relationships among a variable number of adult females. Numbers of females within a pride depend on the size of the pride territory and the amount of resident prey. Females in a pride are not always together as females often break off as solitaries or in groups, especially when a subset has similar-aged cubs.

Male lions born into a pride become nomadic at about two or three years old, depending on a number of factors such as tolerance of the adult females, pride takeover by a new group of males, and number of males in an age group. Such nomadic movements are tentative at first, with young males making short-range movements before returning to the pride territory. Later, movements become more extensive although studies have shown that males tend to settle within a distance of about two pride territories from where they were born.

Depending on the level of competition from other male coalitions, male lions usually stay with a pride for two or three years. One of the most common behaviours attributed to male lions when they take over a pride is that they kill all the cubs belonging to previous males, a behaviour called infanticide. However, studies have shown that infanticide is not as common as believed. Females pregnant at the time of takeover almost always have their cubs accepted by the incoming males. Females with young cubs in a den can also have their offspring accepted as “theirs” by new males. Genetic studies in Etosha National Park in Namibia have shown that 57 per cent of prides contained cubs not sired by the pride males, and out of those four out of 10 cubs had a father not the pride male.

Where can I see lions in the wild

Lions can still be seen in a variety of protected areas in countries in eastern and southern Africa. These destinations are well known and well described, like the Masai Mara, Samburu and Amboseli in Kenya, the Serengeti and Ngorongoro Crater in Tanzania, Hwange in Zimbabwe, Kruger in South Africa, Moremi and Chobe in Botswana and Etosha in Namibia. Of these, the top three places are:

 1. Botswana
Seeing lions in Botswana is probably easiest during the dry season when prey animals concentrate around remaining sources of water – meaning from May to November. For the most reliable locations, head to Duba Plains, Savuti, Chief’s Island and Linyanti.

2. Kenya
Kenya has long been known for lions, though numbers are decreasing. Head for the Masai Mara area, especially the community conservancies, but also the Musiara Swamp where BBC’s Big Cat Diaries were filmed; Amboseli National Park and surrounding conservancies, and Buffalo Springs/Samburu.  Visit from September to March.

3. Tanzania
Tanzania still has Africa’s largest overall lion population and is home to the longest-running lion research programme started by Dr George Schaller in the 1960s in the Serengeti. Visit the Serengeti National Park, the Ngorongoro Crater, and Lake Manyara National Park, and plan your visits between June and October.
There are other places off the lion-viewing beaten track, and those areas deserve more ecotourism attention to inject vitality into their nationally protected areas. These include:

1. The Gambela Park in Ethiopia
A little known treasure located in southwestern Ethiopia, this reserve is home a great diversity of wildlife including over 100,000 white-eared kob and over 400 species of birds. Amazing scenery and very low tourist numbers make this reserve one of the most exclusive safari experiences available on the market today. The reserve is threatened by agricultural schemes and could greatly benefit from a well-deserved influx of ecotourists. The Ethiopian lions could well be very special – recent genetic analysis of lions in the Addis Ababa zoo suggest substantial differences from Kenyan lions for example.
Best times to visit are during the dry season from November to April.

2. Benoue National Park, Cameroon
A UNESCO Biosphere Reserve, Benoue is located in north-central Cameroon. The park consists of savannah, open woodlands and riverine woodland. The National Park is home to one of the largest extant populations of the endangered western African lions, as well as western African elephants and buffalos.
Best times to visit are in the dry season from October to April.
Lion reproduction

Female lions in the wild generally have their first litter when they are about four years old. However, such first time mothers have a poor success rate of surviving cubs – females aged six to nine do better. Overall, in a long-term study of 20 lion females in the Okavango Delta of Botswana, the largest category of adult females died without offspring. The average lifetime reproductive output of females was two cubs, and the best achievement by any female was six cubs over her lifetime.

In the wild, a lion cub has a 60 per cent or higher chance of dying before it reaches the age of two. Such high mortality stems from a number of causes including predation by leopards and hyenas, failure to thrive, abandonment, dying of injuries inflicted by other lions in the pride, not being able to keep up with long-distance pride movements, etc. Survival after age two is better, but becomes difficult again for dispersing young males.

Another difficulty that wild lions have is that they are a semi-social cat species with a reproductive system more suited to solitary cats. All cats are induced ovulators, meaning they need the stimulus of mating before the eggs are presented to be fertilised. The ovulatory cycle in cats requires an initial burst of oestrogen to prime the system, and that works well among solitary species – the male does not usually detect a female in oestrus immediately. With male lions usually in close proximity to females, they might begin mating too early. As mating among lions takes place every 20 minutes or so over three to four days, a male might soon be depleted of any significant sperm count before the female reaches the appropriate stage in her cycle. As a result, most lion matings by a single male do not end in pregnancy – studies in the Okavango indicate that if only one male was present, only 18 per cent of matings resulted in pregnancy. If two or more males mated with the female, pregnancy significantly increased to 73 per cent. 

What threats do lions face?

With current lion population numbers standing at a minimum of 15,000 today versus a possible number of 200,000 only 50 years ago, a number of causative factors have been proposed for this precipitous decline. Most immediately involved are perhaps the intertwined reasons of loss of habitat, loss of wild prey, and conflict with humans/livestock that have occurred as a consequence of the fast-growing human population in Africa. Human populations have not only taken over huge swathes of previously available lion territory, but they have also invaded formerly protected areas, the supposed “safe-houses” for lions. Unless national parks derive income from tourism to provide their needed maintenance budgets, they are soon invaded by cattle and chipped away at by agriculture.

Another major threat to lions are diseases, especially those recently introduced to Africa. In 1994, over 1,000 lions died in a major canine distemper epidemic in the Serengeti and surrounding regions that has subsequently abated to some extent but remains a threat. More and more lions in Kruger National Park are infected with bovine tuberculosis, introduced via domestic cattle to buffalos and subsequently to lions. Lions are particularly vulnerable to infection, as populations can be infected by feline immunodeficiency virus (FIV) at a rate exceeding 95 per cent.

Long dismissed as a threat, more recent information indicates that infection has a significant and cumulative effect on the competence of lion immune systems. At least three of the five major lion populations in existence today (Kruger, Botswana/Hwange and Serengeti/Mara) are thus highly vulnerable to secondary diseases due to eroded immune systems. Experimental studies among domestic cats indicate that FIV is directly implicated in stillbirths, early deaths among kittens, and reduced survivorship among adult cats – a similar suite of problems observed among lions. FIV has probably been among lions for many thousands of years, but lions exposed to newly introduce
d diseases and lack of proper nutrition are additive factors to the consequences of infection.

Despite the precipitous decline in lion numbers, many populations are still subjected to offtake by trophy hunters. Notwithstanding claims that such hunting is sustainable, it has become increasingly apparent that trophy hunting of a small segment of the overall population (adult males) has had significantly negative consequences on lion numbers. Not least because the perturbations caused by selective removal of males within the context of a socially complex species such as lions will have significant effects on reproduction.

Lion viewing guidelines

Lions when relaxed are eminently approachable by vehicles. Remember, however, that such approaches are significantly weighed by the lions’ past experiences. Many guides, eager to give their clients a “close up” experience for photographs and memories, approach lions too closely. When they are resting they might tolerate a close approach, but they do have a limit. If lions walk off, they will not readily tolerate additional attempts to get close. As a general rule, always be respectful of wildlife – and that means keeping a considerate distance and keeping all noise levels down.

If you should encounter lions walking or hunting, such respect should be increased. Lions often detect their prey by hearing, and vehicle noise will impede this. NEVER use spotlights at dusk or night when following moving lions. Their hunting success rate is low enough without the added handicap of noisy vehicles with lights on their tails.


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