Lion

Deirdre Jackson

• Publisher: Reaktion Books
• Year: 2010
• ISBN:
  9781861897350
• Topic: Biological Sciences
• Subtopic: Science General
• Index: Biological Sciences

Although the lion is not the largest, fastest or most lethal animal, its position as king of beasts has rarely been challenged. Since Palaeolithic times, lions have fascinated people, and due to its gallant mane, knowing eyes, and distinctive roar, the animal continues to beguile us today. In Lion, Deirdre Jackson paints a fresh portrait of this regal beast, drawing on folktales, the latest scientific research, and even lion-tamers' memoirs, as well as other little-known sources to tell the story of lions famous and anonymous, familiar and surprising.

Majestic, noble, brave—the lion is an animal that has occupied a great place in the human imagination, inspiring countless myths, lore and legends. As well, this creative relationship has abounded in visual culture—painted on wood and canvas, chiseled in stone, hammered in metal, and tucked between the pages of medieval manuscripts, lions have often represented divinity, dignity, and danger.

In Lion Jackson summarizes the latest findings of field biologists and offers in-depth analyses of works of art, literature, oral traditions, plays, and films. She is a peerless guide on a memorable visual and cultural safari.

1 Lions at Large

Is it possible for us to comprehend the lion, an animal whose way of life differs so dramatically from our own? Is it true, as Ludwig Wittgenstein suggested, that ‘if a lion could talk, we could not understand him’?¹ Our knowledge may never be complete, but this has not deterred wildlife biologists from focusing their binoculars, cameras and microscopes on the great cat, enduring, in their quest for knowledge, extremes of temperature, motor vehicle failures, gastrointestinal parasites, tropical diseases and potentially fatal encounters with their subjects. Several field biologists have almost died of boredom watching lions lying in a languid heap for hours, only to yawn listlessly, roll over and go back to sleep.

Surprisingly little was known of the lion’s behaviour and social organization until the late 1960s when George Schaller, the eminent biologist and conservationist, embarked on his research on lions in the Serengeti, Tanzania. He began his field-work in the summer of 1966, and completed it in September 1969, two months after Neil Armstrong set foot on the moon. Schaller’s research – a giant leap for lions – was widely disseminated and set the standard in the field. Consequently, many people came to believe that the behaviour of the Serengeti lion was representative of the species as a whole, although Schaller had cautioned against this and anticipated that detailed studies of the animal in other habitats might yield different results.²

Since then, lions in Tanzania, Kenya, Uganda, South Africa, Namibia, Botswana, Zimbabwe, Zambia and India have come under scrutiny, but many questions about these extraordinary cats remain unanswered. Lions are idiosyncratic animals that exhibit a remarkably wide range of behaviours. Evidently, they do not read field guides or conform to stereotype. In Botswana, for example, lions have been known to feast on elephants, but prides elsewhere pass on the pachyderm. Animals of both sexes mix in African prides, but males in western India – the sole population left on that subcontinent – rarely consort with females.³ While most young males of the Tanzanian plains leave their natal prides and strike out on their own, those in Kruger National Park, South Africa, rarely stray far from home.⁴ Even within a given habitat, the animals can deviate from one another in actions and appearance. Like the proverbial blind men describing an elephant, field biologists may reach different but equally valid conclusions.

How can we explain variability in lions? The answer lies in the past. Lions are not easily typecast because, over millennia, they have acquired so many different strategies for survival. As we have seen, lions spread from Africa across the globe in a feline diaspora and learned to thrive in various climates and habitats. By the first century, although the lion’s range had contracted dramatically, lions could still be found in Africa, Asia and the Near East. Flexibility held the key to survival: populations of lions learned to adapt to prevailing conditions. Consummate killers, they had no natural enemies – people presented the only threat to their existence. But lions are far more than killing machines and to appreciate their complexity it is necessary to consider both their morphology (physical structure) and behaviour.

Unlike some mammals – cows and sheep, for instance – which are more impassive than poker players, lions have a wide range of facial expressions. A lion with its ears erect, eyes closed and drooping lips is a contented cat. A lion with its ears back, eyes narrowed and teeth bared is a furious feline, best observed from a distance. Ear position is of vital importance to interlion communication, and the distinctive black markings on the backs of the ears probably help lions detect one another when they are lying camouflaged in the long grass.

Every lion has a unique pattern of dark spots whose number, position and size do not change over the course of its lifetime. These are located above and below the top row of whiskers. In the early 1970s, while conducting research in Nairobi National Park, Judith Rudnai noticed the trait and realized that she could distinguish one lion from another simply by analysing its muzzle. Scars and injuries had previously been used to single out individuals, but these are not foolproof signs. Wounds can heal, two or more lions can experience similar injuries, and cubs may not have suffered wear and tear.⁵ The whisker-spot method, a reliable and non-invasive way of identifying individual lions, has been employed by scientists ever since.

Researchers and tourists alike devote large amounts of time, money and effort to view lions in the wild, but animals habituated to humans generally ignore their admirers. As noted by Schaller, ‘lions seldom focus for long on another individual. When they do, their eyes have that peculiar detached gaze which gives the human observer the feeling that the lion is looking around and through him rather than at him.’⁶ The detached expression is not, of course, due to myopia; lions have excellent eyesight and are also equipped with night vision.

The lion’s ability to see in the dark is due to its large pupils and to a light-reflecting layer called the tapetum lucidum (bright tapestry), located behind the retina. In photographs taken at night the eyes of a lion may seem to glow, as if illuminated by an internal light source. The tapetum lucidum, which bounces light back through the retina, is responsible for this effect, transforming an ordinary animal into the possessed protagonist of a Hollywood horror flick. The oval pupils of the lion contract to a pinpoint, rather than a slit, when subjected to sources of strong light, a feature that distinguishes cats in the Pantherinae subfamily (including lions, tigers, jaguars and leopards), from most of their smaller cousins in the Felinae (a much larger and more diversified group, including lynx, cheetahs and domestic cats).

Cats in the Pantherinae subfamily also have supple larynges (voice boxes), featuring an elastic ligament, the epihyoideum, enabling them to roar. Male lions start to roar after their first birthdays and females a few months later, but they do not roar convincingly until the age of two and a half. The sound, a series of plaintive moans building up into a thunderous surge, is one of nature’s most compelling, but it is difficult to describe, let alone imitate, and phonetic interpretations leave a lot to be desired. Lions tend to stand while roaring, but they can roar from any position, even at full gallop. Regrettably, domestic house cats are unable to do likewise because their hyoids are inflexible, but they are compensated by an ability to produce a proper purr, a sound that lions cannot achieve. Lions make a purring sound, but it is a half-purr at best, uttered only on exhalation.⁷

Lions roar in response to other lions, to exert their territorial rights and warn intruders to retreat, to enable pride mates to locate them, to intimidate rivals and to defuse tension. If every skirmish between male lions were to escalate into a raging battle the animals could suffer lethal wounds. By roaring instead males are able to signal their dominance and avoid incurring injuries. Communal roaring, behaviour unique to lions, is exhibited by both wild and captive animals, and is believed to strengthen social bonds. Regardless of its precise purpose, roaring can be infectious; other lions, eager to contribute to the tumult, will often join a soloist, forming mixed- or single-sex choirs. Roaring at the break of day, lions form a memorable dawn chorus.

It was recently reported that a lion, incarcerated in a zoo in Baku, Azerbaijan, repeatedly roared, ‘Allah’. Supporters of the miracle, who filmed the sequence, saw it as a sign of the creature’s devotion to his Creator, but detractors (both Muslim and non-Muslim) pointed out the similarity between the sound emitted by the Arabic-speaking lion of Baku and the standard roar articulated by a lion in the wild.

Stifling a roar can sometimes be more beneficial than expressing it, and lions know when to keep quiet. Studies have shown that nomadic lions, while intruding on the closely patrolled territories of established prides, do not advertise their presence by roaring. In addition, in places where they have been harassed and killed by humans, lions have learned to stay silent to avoid unwanted attention. Furthermore, lions never roar while pursuing their prey.

Playback experiments have proved that lions are able to determine the sex of a roaring lion, to distinguish pride mates from strangers, and to recognize individual voices. When females heard recordings of the roars of their male pride mates, for example, they remained calm, but they reacted with alarm to the roars of foreign males. Single lionesses, played tape recordings of three other females roaring in unison, were generally reluctant to approach, but they were much more willing to advance when played a tape of a lone lioness, because they no longer felt outnumbered. These and similar experiments not only shed light on how lions interpret auditory signals, but also suggest that the cats can count.⁸ But lions are not mathematical geniuses. Females, for instance, do not use counting skills to keep track of their off-spring, and will often abandon a cub or two, inadvertently or otherwise, without acknowledging the loss.

More gregarious than any other great cat, the lion is the only member of Felidae to live in groups and to exhibit marked sexual dimorphism (differences in the form of males and females of the same species). Having inherited territories from their female ancestors, lionesses form the core of the pride, hunting, feeding and raising their young together. Because there is no dominance hierarchy there are no divas or lion queens. A female lion is linked to her birthplace in a fundamental way, and her attachment to her home range never leaves her – she may even prefer to give birth on the very ‘spot where she herself was born’.⁹ Although lions are more social than any other cats, a pride of lions is not an undifferentiated tawny mass; ‘pride members often seem eager to assert their individuality, and the entire pride is very rarely seen together’.¹⁰ If there are too many females in a pride, young ones are likely to be driven away; they may try to stay close to home to establish new prides or they may adopt a peripatetic lifestyle, living alone or teaming up with another lioness.

Pride territories range from 30 to 400 sq km, providing lions with plenty of room to manoeuvre. Males who patrol the home range create a stable social order in which females can raise their young and hunt – a division of labour that enables all members of the pride to flourish. Roles are not firmly fixed. Just as females will help to defend the territory and sometimes kill intruders, male lions make significant contributions by slaughtering big but relatively slow-moving animals, such as buffalo (Syncerus caffer), weighing up to four times their own size. Describing a buffalo kill, field biologist Craig Packer writes:

I once watched a group of females lead their husbands to a buffalo, then stand erect and literally point at the prey: There you are, dears; you can do something useful around the house for once. The males went dutifully forward, hopped on the buffalo’s back, rode it like a bucking bronco, and finally pulled it down. Meanwhile, the females stood perfectly still, cheering the males on from a safe distance.¹¹

There is no disputing a male lion’s ability to hunt, and it is a skill he must rely on during frequent solitary periods or time spent solely in male company without a female cheerleader in sight. Because mature males weigh over 180 kg (400 lb) (they are 20 to 50 per cent heavier than females), they are also more likely to intimidate scavengers, including hyenas that congregate at kill sites.

Lifelong attachments between pride mates are strengthened by displays of recognition and affection. Male lions generally bond with males, and females with females, rubbing heads, licking, lying back to back, entangling limbs or draping a paw over a neighbour’s shoulder. The lion’s tongue is a rough rasp; it can strip meat from the bones of its prey, but it is equally adept at dislodging ticks and mites. Mothers and adult daughters like to groom each other, and cubs can receive such an enthusiastic licking that they are knocked off their feet. Hair is ingested during grooming sessions, and lions suffer hairballs just like their smaller cousins. Sometimes the hair will mat and mix with mineral salts forming a hard, polished stone known as a bezoar. Believed by some Africans, such as the Akamba of Kenya, to have magical and protective powers, and to confer on the bearer immunity from lion attacks, the stone was once a treasured talisman. Dying lions were said to spit out the object, and hunters would search diligently for it.

Lions often sniff each other’s genitals – a form of greeting that tells them many things, including whether females are ready to mate. Rocks, trees, shrubs and the odd Land Rover are sprayed with urine and the scent from a lion’s anal glands, and these markings contain essential data for both scientists and lions alike. As noted by Bruce Patterson, ‘anyone who has smelled cat urine knows that it contains plenty of volatile components that have the potential for copious biological information’.¹² Lions make a distinctive scraping motion with their hind feet and often urinate on the ground afterwards. By sniffing pugmarks they are able to track one another’s movements and determine whether trespassers have crossed their pride boundaries. When they smell something particularly interesting, like carrion or urine, lions will raise their muzzles, wrinkle their noses, open their mouths and close their eyes – a gesture known as flehmen. This grimace exposes the vomeronasal organ on the roof of the mouth, and enables them to decode scent molecules and pheromones.

Lions’ lives are not the frantic and emotionally charged soap operas portrayed in heavily edited wildlife films. In fact, lions spend 80 per cent of their time doing next to nothing. A lion is an overstuffed armchair, a stalled car, a rock. With his eyes half closed he stares into the distance, thinking lion thoughts, dreaming lion dreams. Flies buzz. A single cloud shaped like a croissant drifts across the sky. There is not an awful lot for a researcher to report. As the indefatigable Schaller monitored the sleepy lions of the Serengeti, Rudnai scrutinized their drowsy counterparts in Nairobi National Park. After measuring intervals between yawns and observing group dynamics, she discovered that a lion can yawn up to five times in twelve minutes, and that yawning is just as contagious in lions as it is in people.¹³ Whether the lions had a soporific effect on either scientist remains unclear.

Disappointingly, lions are not inclined to prolonged courtship rituals. There is no leonine equivalent of the leaping and bowing performed by cranes. Biting, cuffing, snarling and growling are standard behaviour for mating lions. What they lack in grace, however, they make up in stamina – they are capable of mating once every twelve minutes for up to six days. Coitus stimulates ovulation; for every successful pregnancy lions may copulate up to 1,500 times.¹⁴One promiscuous lioness mated so many times that ‘her rump sported a smooth, shiny spot from constant wear’.¹⁵ Given the lion’s robust sex life, it is not surprising that in some parts of Africa sleeping on a lion’s skin...