The Lion

Of all the great cats, the lion has always held a supreme place in man's esteem and imagination. The lion has always been honored by man, crediting the regal beast with attributes he prizes most; nobility, courage, loyalty, combative skills and sexual prowess. This attribution applies to both sexes, for a lioness is a creature of sinuous beauty, but the full-grown male, whose magnificent mane ranging in color from a rich golden brown to a deep blackish-brown, mark him as the veritable monarch of the plains.

The legacy of the lion, King of Beasts, as the model throughout history is demonstrated by its appearance among the earliest drawings made by humans over 15,000 years ago. The myth of the supernatural powers of the lion survives today; by consuming or wearing parts of a lion it is believed that one can revive lost powers, cure illness, attain courage and win immunity from death.

Yet for all its glory, the lion is only the second largest of the big cats, the tiger being the largest. Full grown males displaying a regal and imposing nonchalance, regularly exceeding 400 pounds in the wild, and measuring up to 10 feet in length, from the tip of the tail to the nose. Females, only slightly smaller, range from 275 to 400 pounds and measure up to nine feet in length. Like other members of the Felidae, the lion has a lithe, compact, muscular and deep chested body with a rounded and shortened head bearing prominent whiskers.

Biology

The lions, one of the five species of the genus Panthera, are composed of seven subspecies; the Angolan lion, endangered, Panthera leo bleyenberghi; the Asiatic lion of India's Gir Forest, endangered, Panthera leo persica; the Masai lion, Panthera leo massaieus; the Senegalese lion, endangered, Panthera leo senegalensis; the Transvaal lion, endangered, Panthera leo krugeri; the Barbary lion, extinct, Panthera leo leo; the Cape lion, extinct, Panthera leo leo; and the Cape lion, extinct, Panthera leo malanochiata.

The lion's mane, the only instance of intersexual variation among the felids, usually begins a rapid develop, from a small tawny ruff at the age of two years old, into a massive thickened mass of hair often extending from the neck area to the underside of the abdomen. These morphological changes are indicative of the onset of sexual maturity, and hasten the ejection of these adolescent males from the pride by the dominate males. In the context of pride structure and defence, the mane has clear functions and advantages. The male's mane increases his apparent size, serves as an visual indication of gender from long distances and adds to his aura during his strutting displays in front females. The mane may also be a visual basis for a potential opponent to guage the vigor and age of the male and functions as well to protect its owner against the claws and teeth of an opponent should fighting occur.

Male Dominance

The lion is the most social of all felids, setting them apart from all other great cats. Life in the pride is, for the most part, a tactile experience with a great deal of social grooming and rubbing of heads and bodies. The pride, which varies in size from as few as four individuals to as many as 40, is composed of a group of often related lionesses, their offspring and one to six nomadic males who have fought their way to dominance. These newly "installed" dominate males generally kill, and often consume, all of the cubs belonging to the previous males, presumably to enhance the chances of survival for any cubs they sire. Within a week or two, the lionesses respond to the absence of cubs by coming into season and maiting with the dominate male(s). The members of a lion pride are sometimes scattered in several groups throughout the pride's range.

The size of these groups, sometimes called "companionships" or "subprides" is influenced by a number of ecological and social factors including the availibility and size of prey species. A pride will range over an area of between 10 and 175 square miles, often overlapping the range of neighboring prides, depending mostly on the movement and abundance of prey. The core of the female oriented pride are the lionesses, who's association is based on matriarchal continuity. Lionesses not only assume the responsibility of hunting for the group but also care for the offspring, who suckle indiscriminately from any or all of the nursing females in the pride. Except for fighting over the right to feed at a kill, where male lions are the first to eat with the females and cubs initially sidelined, pride members get along harmoniously, displaying warm signs of affection by rubbing heads and cheeks when the greet one another and licking one anothers faces after feeding. The resident male lion(s) breeds with the females when they are in season, and contributes to the pride by maintaining stability by protecting the entire group from dangerous intruders.

Hunting Methods

In characterizing hunting techniques within the pride, there seems to be some correlation between prey availability and/or prey size and the number of lionesses willing to participate in any hunting effort. If the prey species is small and a single lioness is capable of making a kill without the help of others, pride members are often content to wait and perhaps gain a free meal. If the prey species is large enough to feed the entire pride, and a joint hunt is more likely to succeed, cooperative hunting appears to benefit all participants. Although hunting techniques vary from single lionesses to group efforts, depending on local conditions and the willingness of pride members exert themselves and risk injury, the pride structure is highly effective when it comes to rearing young.

After a lioness leads her cubs from their hidden birthing dens, about one month after birth, the effectiveness of the pride structure becomes evident. Staying close to their mother at first, the cubs soon begin to explore their new surroundings and the littermates of other pride members. Lionesses' with cubs form sub-pride groupings, referred to as creches, to maximize the care, protection and food resources available for successfully rearing young. Within these cheches cubs are nursed by all members of the group although milk distribution depends, in large part, on a pattern of surplus production by individual lionesses and a preference for kinship. Generosity among lionesses is somewhat indiferent, with respect to milk, and crechemates tend to feed from lionesses who have an excess supply, generally the result of a small litter . This communal feeding is often accomplished after a lioness has fallen asleep and not able to prevent any other than her own offspring from nursing.

Lions are not particularly efficient hunters, successfully capturing prey consisting of medium sized ungulates including zebras, wilderbeasts and antelope in only 20 to 30% of their attempts. They are referred to as "opportunistic" hunters, eating whatever they can catch for themselves or steal from other predators. They are not well adapted for leaping or reaching particularly high speeds, nor are they capable of running for long distances. In general, if the lion is not successful within a few hundred meters, they give up the chase and the prey escapes. Two important causes of hunting failure relate to a fault in the actual stalking of the prey and in the execution of the characteristic lion charge. Lions do not hunt by scent, although their sense of smell is excellent, they often approach prey from an upwind location thereby alerting the prey and ending the hunt. Secondly, the lion's charge is generally launched directly at its quary and it rarely alters the path of the attack, as do other felids.

Generally speaking, if a lion misses it's quary on the initial charge, it does not give pursuit, but quits and looks for new quary. Scavenging is also an important source of food for lions, with food stolen from other predators and or carion often making up 10 to 15 percent of their total food intake. Field observations reveal that lions spend a great deal of time looking for circling vultures and listening for the calls of hyenas, enabling them to locate downed prey. When prey and other predators are plentiful, lions may get close to half of their food by scavenging. The sharp eyesight of antelope, wilderbeast and zebras makes stalking by day, on the open plain, extremely difficult. During an early morning stalk, especially when several lions are taking part, other animals of the grasslands including the intended victims know that the lions are hunting. Unless a lion is in sight, in which case all eyes will be fixed on it, the prey will be glancing around nervously. Early flight is not always the best method of escape for the quary because they do not know where all the lions are located nor do they know how or where they will strike. Evolution seems to have dictated that they wait until the lions commit themselves to the attack, acting prematurely, and in confusion, are apparently less effective in escaping these predators.

Approaching the prey at speeds of up to 35 miles per hour, the lioness uses its body weight, large paws and claws to bowl all but the largest prey over, and secure it to the ground. Killing is normally accomplished in one of three ways; a nape of the neck bite for small prey that severs the spinal chord; a throat bite for larger prey that kills by strangulation; and a muzzle bite that also suffocates the quary. Although lions hunt during all times of the day, nocturnal hunts are generally more successful. Therefore, lions in many area of their range, prefer to hunt under the cover of darkness where the light gathering adaptations of the felid eyes casts a distinct advantage to the predators. As darkness approaches, the lionesses silently move out in lines to locate prey, circling around and behind a herd they pick out a startled victim and dispatch it with a bite to the neck or throat.

Feeding Habits

Interactions and pecking order among pride members at a kill is highly developed and lions rarely eat in peace. The uncertain temper and tremendous strength of the grown males, feeding at kills, make confrontation a risky business. Therefore, the mature males, if present, always eat first and rerely tolerate females in the initial feeding when the full pride is assembled. When they have satisfied themselves, they retire in the immediate vicinity to relax, and the females come forward with constant bickering and fighting among themselves. Interestingly, most of the facial wounds, and the injuries represented by the scars on the lions heads, are received during squabbles at kills. Juveniles and cubs are the last to feed and are frequently left out altogether. A sad and all but inexplicable fact of lion life, given the generally social nature of the pride, is that the leading cause of cub death is starvation. Cubs are apparently readily expendable, in the interest of the pride during periods of food shortage, and are later easily replaced when food supply is increased.

Intra Pride Relationships

Lions have an instinctual and ingenious method of limiting serious injury during potentially violent confrontations. The weaker members of the pride, especially cubs, adopt a fawning or cringing position and, by crouching or lying submissively on its back, deliberately exposes itself in such a fashion as to make killing as easy as possible for the superior lion. The aggressor, recognizing the appeasement gesture and that it is no longer threatened, breaks off the attack and returns to other matters. Disputes between mature males are not generally governed by the normal etiquit extended to other pride members. These confrontations can be extremely violent or fatal encounters, where submissive gestures and other body posturing can result in severe injury or death to the weaker individual.

Female lions generally come into estrous by the age of 2 1/2 to 3 years and continue to have litters every two years, if an adequate food supply exists. Hormonal changes in the lioness are detected by the dominate male(s) through changes in the odor of the lionesses urine. When smelling the urine of an estrous lioness, the male lion makes a characteristic facial grimace, known as "flehmen." The male lion raises his head, swills the females odorous urine in his mouth, and inhales deeply, drawing back his lips in a curl exposing his teeth and gums. Male lions become extremely protective of their mates while they are breeding, often leaving the immediate vicinity of the pride. As a prelude to maiting, generally initiated by the lioness, body posturing including rubbing and nudging, rolling over on her back and emitting a low guttural moaning stimulate the male to initiate intercourse. When she has his full attention, the lioness assumes a crouched maiting posture with the base of her tail slightly elevated. The male mounts and copulation lasts from 5 to 20 seconds, during which time the male bites the neck of the lioness and produces any number of facial expressions and sounds. At this time lions mate frequently, as often as every 20 minutes or half hour, for several (3-7) days, the estrous cycle lasting from one day to three weeks. Following each copulation, the lioness rolls over onto her back and lays in this inverted position for a minute or two, perhaps enhancing the chances for conception. Maiting lions do not usually show any interest in hunting or eating, and are not generally viewed as a threat by prey species. Studies have shown that approximately one cub reaches sexual maturity for every 2,500 copulations.

Lion Cubs

As the end of the 105 day gestation period approaches, the lioness leaves the pride to seek out a sheltered isolated area to give birth to her litter of two to four cubs, as a matter of safety to the cubs. The cubs are born blind, weighing from three to four pounds, with a soft wooly greyish-yellow or pale brown coat variously marked with spots or stripes. The grey blue eyes of the cubs turn into the amber color of the adult's eyes at between two and three months old. The lioness rarely leaves her young cubs. When she does, to hunt or drink, she risks loosing them to other predators like cheetahs, leopards, jackals or hyenas. Under most circumstances, cub mortality is high, with more male deaths than females, perhaps because males are more active and more likely to turn up lost or in some fatal predicament. This disproportionate infant mortality results in the male-female imbalance observed later in life. As the cubs grow, the lioness begins to devote much of her time retrieving her adventurous youngsters who have ventured from the safety of the hiding place. At four to six weeks of age, when they able to walk fairly well, the lioness leads the cubs from their hiding place and they are introduced to the pride. Adult lions are fond of cubs, and the introduction of new ones is an occasion for a good deal of solicitous examination. The new cubs settle very quickly, and in a few days have become full members of the pride.

A day in the life of an exuberant lion cub might be described as a gregarious melee of social interaction. The curiosity of lion cubs is boundless. Cubs wrestle for hours, chase one another and anything that moves, crawl on and spar with their mother, climb trees or any other structure that is available and even interact with adult males, with inevitably mixed results. In nature's plan, all the playful tussling has a purpose; the young cats are, in part, exercising to develop their muscles and to sharpen their reflexes and, in part, working out the fighting techniques that they will need as adults to defend their territories, prides and food against rivals. In large prides, cubs will vary widely in age, ensuring that the youngest learn early the rules of survival. Cubs playing help develop the techniques that adult lions employ in hunting: stalking, rushing, crashing, chasing and pulling one another down from behind; all as will be used someday against prey. Cubs begin to accompany their mother on hunting expeditions when they are only about three to four months old although they will not kill their own prey until they are about two years old.

Male lion cubs begin to show visual signs of sexual maturity by the age of twelve months (mane), but only begin to exhibit true adult male behaviors about the age of 30 months. The dominate male lions of the pride slowly recognize their behavior and display hightened agressiveness towards the adolescents, until one day they drive them from the pride. These adolescent males tend to remain on the periphery of the pride for a few days before striking out completely on their own. Ousted males often form "coalitions," typically consisting of brothers and/or cousins that have been reared in the same nursery groups. Other groups consist of non-relatives that teamed up during their solitary nomadic wandering. A male lion's reproductive success depends directly on how well his coalition can withstand challenges from outside groups of other males. Male lions display their greatest capacity for teamwork when challenged with ousting other nomadic males intent on dominating the pride. The typical duration for which a strong coalition maintains control over a pride is two to three years, during which time they father all offspring born in the pride.

Sleeping

Lions devote an inordinate amount of time sleeping, especially at midday on the plains, once having found the shadow of an acacia tree. After consuming a full meal, the pride may spend the next 24 hours just resting and digesting their meal. Being extremely social, lions lay about completely intertwined with heads and legs lying over and around adjacent individuals. When one shifts positions, the chain reaction affect many of it resting partners.

Lions produce a unique vocalization which may be produced by all adult members of the pride. Roaring is the loudest sound uttered by any cat, recorded as loud as 114 decibels, and can be heard from a distance of up to five miles. Generally, roaring is performed between dusk and dawn, often in response to the roars of other neighboring prides. Roaring links the members of a pride, as well as strengthening their claim to territory and deterring possible trespassing by neighboring prides. This system of territorial behavior appears to be an effective mechanism for reducing disruptive competition for food. The sound can be produced when the lion is lying on the ground, but most often in a standing position, its head lowered and back arched; the face not relaxed but taut; the neck stretched forward, with the muscles of the throat tight; eyes usually open; mouth only partly ajar; and the nose only slightly wrinkled. The entire body is taut with the abdominal muscles rapidly tightened and contracted to exhale to air required to produce the roar.

Man Eatings

Lions become man-eaters less often than tigers, but when they do they are bolder and more aggressive in their pursuit of humans. A man-eating lion often hunts at night and prowls the perimeter of villages looking for victims. One lion in Uganda killed 84 people before it was killed in the early 1920's. During the construction of the Tsavo River bridge in Kenya, a number of lions were reported to be man-eaters and terrifying stories of their nighttime raids kept many workers within the confines of the living compounds.

An isolated group of white lions has been identified on the plains and scrublands of South Africa in North Transvaal, Timbavati Game Reserve and North Kruger National Park. The light coloration is due to a recessive gene much like that which produces the white bengal tigers. The pelage color is ivory to cream with light golden eyes, and therefore they are not albinos. Cubs are typically almost pure white at birth, but change to the ivory/cream coloration as they mature. In the United States, the Philadelphia Zoo and preformers Siegfried & Roy, at the Mirage Hotel in Las Vegas, maintain and breed white lions.

Diseases

Recently, canine distemper virus (CDV) has been identified in the lions of the Serengeti Park. The virus appears to have been transmitted from the domestic dogs living adjacent to the park. In 1978, CDV is thought to have caused several fatal epidemics in canids within the Serengeti-Mara ecosystem of East Africa, affecting silver-backed jackals (Canis mesomelas) and bat-eared foxes (Otocyon megalotis). In 1991 CDV was identified in the African wild dogs (Lycaon pictus) in the ecosystem. The large, closely monitored Serengeti lion population was not affected in either of these epidemics. However, an epidemic caused by a morbillivirus, closely related to the CDV, emerged abruptly in the lion population of the Serengeti National Park, Tanzania, in early 1994. This epidemic manifested itself in a fatal neurological disease characterized by grand mal seizures and myoclonus; the lions that died had encephalitis and pneumonia. By August 1994, eighty-five percent of the Serengeti lion population had anti-CDV antibodies, and the epidemic spread north to lions in the Maasai Mara National reserve, Kenya, and uncounted hyenas, bat-eared foxes, and leopards were also affected. There is now an effort to "ring-vaccinate" around the park, using a new vaccine. This inoculation is being carried out by Cambridge (UK) veterinary graduate student Sarah Cleaveland, who vaccinates the dogs in the area every six months. This will hopefully create an area around the Serengeti Park with fewer infected animals, reducing the possibility of transmission to the lions. It was decided to ring-vaccinate the dogs, rather than the lions themselves, due to the (small) risk of the vaccine virulently infecting a proportion of lions being vaccinated, and also to the difficulty of actually vaccinating lions in the wild. The majority of infections are asymptomatic, because the immune system of most victims neutralizes the virus before it spreads to the central nervous system. This selection of animals, with immune systems fast enough to respond, would explain why only 1,000 out of 3,000 Serengeti lions died, and the others weren't observed to be affect


 

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