LION, Panthera leo

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WHAT IT IS

Call of the African Wild, King of African Carnivores. Low hut large and powerful, especially males. Male wt 416 lb. (189 kg) (record 572 lb. {260 kg} Namibia), ht 48 in (120 cm); Female wt 277 lb. (126 kg), ht 44 in (110 cm). Coat short except for tail tuft and male's mane appearing third year, maximum development at 5. Color: tawny with white underparts often faintly spotted (especially in East Africa); black tail tuft, ear backs, and lips; mane individually variable, from blond to black; cubs woolly with grayish, spotted coats, changing to adult coat by 3 months. Teats: 4.

WHERE IT LIVES

Sub Saharan Africa except deserts and rainforest, wherever medium sized and large herbivores survive. Still the most abundant large predator (after spotted hyena) in savanna and plains ecosystems. Formerly ranged Asia and Pan Africa, but exterminated in North and South Africa by early this century (except for Kruger and Kalahari Gemsbok NPs and recently reintroduced in some Transvaal and Natal reserves).

GOOD PLACES TO SEE IT

Just about all the major NPs. The problem in more popular ones is finding lions unaccompanied by vehicles.

ECOLOGY

Savanna and plains habitats with greatest variety and biomass of hoofed mammals carry up to 1 lion/3 square mile (12/100 square kilometer). Where prey density is very low, as in Miombo Woodland Zone or Sahel, there may be only 1 lion/50 to 100 square mile. Commonest ungulates from impala to wildebeest and zebra in size are main prey. Different prides have different preferences and traditions. Some, hunting in groups usually including males, regularly kill buffalos, including biggest, oldest bulls; even bull giraffes are occasionally taken (caught lying down). Variety of smaller game also taken by hungry or curious lions, including rodents, birds, turtles, lizards, even fish, and ostrich eggs (by the few with knack of opening them). Another important source of food is scavenging. Lions respond to rallying cries of hyenas as readily as hyenas themselves and by day are guided to carcasses by descending vultures.

ACTIVITY

While prey is plentiful, lions spend 20 hours out of 24 conserving energy, becoming active in late afternoon when mothers retrieve, suckle, and socialize with young cubs and one another; hunt most actively early and late at night, carrying over for a couple hours after daybreak. But lions become active any time, day or night, hungry or gorged, that easy opportunities to catch prey present themselves.

SOCIAL/MATING SYSTEM

THE FEMALE PRIDE

The basic units of lion society are prides of related females, each pride residing in a traditional home range/territory. Male offspring have to leave by 2.5 years. Resident adult males are immigrants that have gained custody of a pride range in competition with other males.
The number of adult females in a pride is adjusted to seasons of minimum prey availability and tends to be consistent over time. Surplus females have to disperse; if the membership falls below capacity, subadult nomads are accepted in the absence of recruits from within the pride. Home range size also depends on prey density, being as small as 8 to over 154 square mile (20 400 square kilometer).
In Serengeti and Kruger NP, a typical pride numbers about 13. The aver age composition of 14 prides totaling 181 lions was 1.7 (1-4) adult males, 4.5 (2-9) adult females, 3.8 subadults, and 2.8 juveniles and yearlings. Large prides, which can include up to 40 lions, may never assemble in one place. Members come and go unpredictably, alone and in groups, typically numbering 3 to 5 lions. There is no rank hierarchy among females and no 2 are likely to be found together more than half the time. But all residents are acquainted and whenever they meet, the lion greeting ceremony reaffirms their social ties. A lion without the self assurance to meet and greet sends a signal that it doesn't belong and is treated as an intruder. Each sex defends the part of the pride range in current use against intruders of the same gender.

MALE COALITIONS

The opportunity to monopolize reproduction of a whole group of females is behind the pronounced sexual dimorphism seen in lions, alone of all cats. The advantages of large size and a showy (also protective) mane have also caused males to become so bulky and conspicuous as to reduce their hunting ability. Reproductive competition is so fierce that males form coalitions to improve their chances. Where lions are plentiful, a single lion has little chance of winning or holding a pride's territory. Once begun, the advantage of competing cooperatively should theoretically lead to bigger and bigger coalitions, ending up in gang warfare; yet coalitions of over 4 males are rare. Large groups have problems, starting with assembling and coordinating all the members. More important, a big coalition destabilizes lion society by taking over different prides then failing to defend them all, with fewer surviving offspring the end result.
Coalition partners are usually related males that left their pride as adolescents and stayed together as nomads until mature and ready to compete. Lone nomads also join forces and can form coalitions as cohesive as sibling teams.
Infanticide is another consequence of severe competition. Prime years for males are 5 to 9; even at 8 they are already losing weight and mane hair; few survive past 10. Having taken over a pride territory, often after a battle royal entailing severe injury and even death for the losers, the victors usually have only 2 years before losing in turn to a younger and stronger or larger coalition. Large coalitions of 4 to 6 males may last up to 4 years; losers don't get a second chance.
Given so little time to propagate their genes, sexual selection rewards males that kill all the suckling young of defeated rivals. When a nursing lioness loses her cubs, she comes into heat within a few weeks. Otherwise, the normal interval between births is 2 years—matching average male tenure. It is thus entirely understandable, though no less gruesome, that the first thing males do after a takeover is kill all the cubs they can catch. Lionesses may fiercely attack them, and in concert can sometimes stop infanticide, but mothers also get killed or have to flee with their offspring.
After a takeover, lionesses often come into heat every few weeks for 4 or 5 months without becoming pregnant. This interval of sexual hyperactivity turns out to be a form of insurance against desertion newcomer males are likelier to become bonded to the lionesses and settle in their territory after months of feverish sexual activity than if the females conceived and stopped cycling the first time around. Birth control also allows time for the biggest and toughest male group to move in and take over. When the sterility period ends, pride females ovulate, conceive, and litter in synchrony. The cubs are then reared communally, improving the chances that a sizeable male coalition will eventually propagate their parents' genes.

FORAGING/PREDATORY BEHAVIOR

Like other cats, except for communal hunts. These usually involve 3 to 8 lionesses moving on a broad front in an attempt to drive quarry into an ambush or block the escape route of ungulates feeding in a cul-de-sac as when wildebeests or zebras graze alongside a river or woodland edge. Males only hunt for themselves when no free lunch is provided by lionesses, hyenas, or other agencies.
Lions hunting in twos and groups have a success rate of c. 30%, compared to only 17 to 19% for lions hunting singly by daylight. But recent studies indicating that single hunters are about as successful as groups at night reopen questions about the primary reason lions became the only sociable cat. Maybe it was to control exclusive hunting grounds and share food with relatives, while protecting it against competitors, including other lions. But lions share food grudgingly; they often fight for places at a kill and prime males take the proverbial "lion's share"- up to 25% of their own weight. If the kill is small, the smallest and weakest lions lose out—hungry mothers won't share even with their own youngsters. High juvenile mortality rates during times of prey scarcity are the result. Nevertheless, when there is enough meat to go around, the whole pride prospers.

REPRODUCTION

Year round but often synchronized within prides— perhaps mainly as a result of male takeovers and infanticide. Typically 3 cubs/litter, after 14 to 15 week gestation; 20 to 30 months between births. Females start breeding at 4, only a year earlier than males.

MATING

For every cub that survives to yearling stage, lions copulate an estimated 3000 times. Only 1 estrus in 5 results in progeny and estrus lasts c. 4 days, during which couples mate 2.2 times/hour. Surprisingly, coalition partners hardly ever fight over mating rights. The first to reach a female in heat becomes her consort—until and unless he has had enough. As partners are usually equals, fighting would impair their ability to withstand takeover attempts.

OFFSPRING AND COMMUNAL CARE

Weighing only 2 to 4.5 lb. (1 2 kg) at birth, lions are helpless as any kitten. Eyes open at 3 to l days, walk at 10 to 15 days, run at 1 month. After 4 to 8 weeks in hiding, mother begins leading cubs to nearby kills. By 7 weeks they keep up with pride. Weaned at 1 to 10 months but remain dependent until 16 months at least. Mothers rarely bring live prey for cubs to practice catching.
Cub survival is highest when reproduction is synchronized, since communal suckling is most equable when there are no bigger cubs to hog the milk.
Juveniles, often left alone for over 24 hours, are vulnerable to other predators that happen on their hiding place. Mothers won't wait for juveniles older than 5 to 7 months; when large prey is scarce and mothers grow thin, they often abandon weakened cubs unable to keep up, especially if there is only one.

RELATIONS WITH OTHER PREDATORS

Either competitive or predatory. Lions kill and often eat all the other carnivores, including leopard and cheetahs (but rarely hyenas see spotted hyena account).

DIFFERENCES IN LION BEHAVIOR


Advertising Presence and Social Status


SCENT MARKING

Mainly in a territorial context.
Urine spraying bushes, high grass, etc. Resident males regularly patrol and spray bushes in currently defended part of pride territory. Females spray only occasionally.
Scuff marking + urinating, by both sexes. Beginning at 2 yr.

CALLING

Roaring: usually standing or crouching but possible from any position and even while running. Roaring grades from soft huh to full throated roars audible 5 mi. Multipurpose behavior: advertise territorial occupancy, contact pride members, strengthen social bonds (roaring in chorus); intimidate rivals during aggressive interactions. male’s roar deeper louder than females, but sex and distance are both hard to judge.

Aggression

Strutting. male dominance display, directed mainly to females,
Head low threat lion keeps head low, forelegs wide apart, shoulders higher than normal, gazing steadily at opponent with eyes and mouth as in typical offensive threat or while snarling (more defensive); ears twisted so that black marks face forward. Unlike threat posture of other cats. If tail is lashed up and down during display and the lion growls or coughs, a charge is imminent.

Sociable Behavior

Greeting ceremony. Lions moan softly and lean on one another so hard that when one is lying, the other often falls on top. Cubs more often rub against adults, and females rub against male-the direction being from weaker to stronger.
Males and female coalition partners lying in contact. Characteristic, another sign of close social bonds.

CALLS

Humming. A sound of contentment, like purring, emitted during affectionate interactions and while cubs nurse.
Puffing, a faint pfff pfff emitted with closed mouth as lions approach each other. Signals peaceable intentions.

Play

Including adult females. females remain playful as adults, whereas males lose the tendency at about 3 yr.

Response to Competitors/Predators

Woofing. Sound made by a startled lion.
Reprinted from "The Safari Companion" by Richard Estes
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