SPOTTED HYENA, Crocuta crocuta

WHAT IT IS

A vaguely bearlike large hyena, the only one with spots. Sturdy build, high shoulders, and long muscular limbs. Male wt 123-138 lb (56-63 kg), ht 32-34 in (79-86 cm); femalewt 147-165 lb (67-75 kg), ht 34-35 in (84-89 cm). Head: massive; broad, rounded ears. Teeth: robust, outer incisors like small canines. Coat: rough, comparatively short. Color: reddish brown to tan, growing lighter and less spotted with age; yearlings gray and heavily spotted; natal coat nearly black and unspotted; tail pompom and nose black regardless of age. Genitals: boneless phallus 5.8 to 7.8 in (14.5-19.5cm), equally developed in both sexes; female differences develop at puberty, when the urethral opening splits and teats enlarge,

WHERE IT LIVES

Everywhere south of the Sahara except rainforests and true desert, up to 13,000 ft (4000 m.). Largely exterminated in South Africa.

GOOD PLACES TO SEE IT

Many parks in East and southern Africa; especially Ngorongoro Crater, Serengeti NP, Tanzania; Masai Mara NR, Kenya; Savuti Pan, Chobe NP, Botswana; Kruger NP, South Africa; Etosha NP, Namibia.

ECOLOGY

The most abundant large carnivore in areas where antelopes and zebras abound. More carnivorous than other hyenas, Crocuta is a formidable predator/scavenger that lives wherever animal resources are adequate. Eats vertebrates of all kinds, especially hoofed mammals; seldom eats invertebrates, fruits, or vegetables. Utilizes carcasses of large vertebrates more efficiently than other carnivores, which waste up to 40% of their kills. Eats everything but rumen contents and horn bosses of the biggest antelopes, even deriving nourishment from mummified carcasses. Bones, horns, hooves, even teeth are digested completely within 24 hours.

ACTIVITY

Radio-collared female, followed in Ngorongoro Crater for 12 days, spent little time foraging and eating, but simply shared kills made by other clan members. She traveled less than 4 mi (10 km) in 24 hours, spending 84% of the time lying down. In Kalahari Gemsbok NP, where hyenas have to work for a living, clan members traveled an average of 16 mi during nocturnal foraging and remained active 31% of a 24-hour day.

SOCIAL/MATING SYSTEM

Lives in large clans and defends territory at high density, but differs from all other social carnivores:

Clan members compete more and cooperate less than most social carnivores.

Females are bigger than males and dominate them.

Females compete for rank and food with one another; even close relatives do not cross-suckle offspring.

Cubs are raised in communal dens, but seldom or never provisioned or guarded by clan members.

Males play no parental role-only a privileged few are permitted anywhere near dens, where even juvenile offspring of high-ranking females dare to bully them.
In Ngorongoro Crater, where a resident population of 20,000 herbivores supports Africa's densest hyena population (approximately 450 adults and their offspring), 7 different clans of 35 to 80 hyenas ferociously defend their sections averaging 12 mi square (30 km square) of the 100 mi square (260 km square) crater floor against their neighbors. At the opposite extreme, clans of 3 to 12 hyenas in subdesert Kalahari Gemsbok NP have ranges of 193 to 772 mi square (500-2000 km square) -much too vast to be defended as territories.
Members of a large clan seldom if ever all assemble at one place. However, a communal den shared by up to 10 females is a social gathering place for them and their dependent and older offspring. Each clan member knows its place in the hierarchy and behaves accordingly whenever clan members meet. Even without behavioral clues, dominant females are usually recognizable as the biggest, fattest hyenas, with swollen udders. Females not only dominate males at kills and other favored sites but also lead clan members on pack hunts, boundary patrols, and into battle.
Female offspring remain in their natal clan; males disperse at around 2 years. Sons of high-ranking females tend to emigrate later and, thanks to the large size and selfconfidence gained as their birthright, are likely to become dominant breeding males in the clans they join. The ensuing reproductive jackpot is what drove females to compete for dominance. But to get there, they not only had to grow bigger than males, they also had to produce male sex hormones to become more aggressive. In the process, both their behavior and anatomy became masculine.

HOW IT MOVES

Ambling walk, rarely trots, lopes tirelessly at 6 mph (10 kph), and can gallop 25 to 31 mph (40-50 kph) for a couple miles at least. Top speed approximately 37 mph (60 kph).

FORAGING/PREDATORY BEHAVIOR

Reputation as skulking scavenger, craven coward, and killer of young and sick animals applies to relations with humans. As scavengers around settlements, hyenas are persistent but remarkably craven when confronted. Similar caution combined with great patience is shown by hyenas awaiting the right moment to finish off sick or crippled large animals. Perfect opportunists, they always take the path of least resistance. Yet Crocuta is also the second-biggest and most formidable African carnivore, capable of running down and killing unaided a bull wildebeest 3 times its own weight.
How does the craven scavenger become transformed into the bold predator? Hunger drives a hyena to run down a large healthy ungulate-but only after failing to get a meal with less effort and risk. Predation also occurs spontaneously-e.g., when a daytime thunderstorm makes the ground slippery and antelopes easier to catch than usual.
Most hunting, like most foraging, is done alone. But clan members sometimes deliberately set off in packs to hunt specific quarry, such as zebras. More often, what look like pack hunts begin as chases by 1 or 2 hyenas that others passed en route join. Whenever 2 or 3 hyenas feed on the same carcass, they begin squabbling; by thus broadcasting the presence of meat, they unwittingly invite all pack members in earshot to the feast-and often enough, lions also get the message. Pitched battles between rival hyena clans are likely when kills are made near borders.

REPRODUCTION

Nonseasonal; 2 young/litter (range 1-4); gestation 4 months. Maturation at 3 years, females later than males.
The female's bizarre reproductive tract, entered and exited through the phallus instead of directly-the vagina entrance being blocked by a false scrotum and testes-makes copulation complicated. When hyenas mate, only the male has an erection-the female's phallus is slack, the prepuce a wrinkled ring around the enlarged urethra. The male has only to slide beneath the female to enter, an operation assisted by the upward angle of the fully erect penis; following which, he straightens into a normal mating stance.
Female dominance imposes unusual diffidence and patience on males. Their obvious fear of female aggression makes courtship an often amusing spectacle.

OFFSPRING AND MATERNAL CARE

Communal denning but no communal care.
Long gestation results in unusually precocious offspring. Newborns weighing 2.2 to 3.6 lb (1-1.6 kg), already have milk incisors and canines, open (but unseeing) eyes, and capability of strong, directed movement on their forelimbs. Two to 6 weeks after whelping inside the entrance of an unused aardvark or other burrow, other transports young to a communal den containing youngsters of different ages. Starting at 6 times/day, suckling is reduced to 3 times/day at 3 months, in bouts lasting up to 1.5 hours! Weaning takes several months beginning at 1 year, marked by frequent tantrums. Cubs of alpha female go to nearby kills and begin eating meat as early as 3 months, 4 to 5 months earlier than rank-and-file offspring. Yearlings tag along on hunts but rarely become competent before 1.5 years.
Beginning only hours after birth, siblings of like sex battle for dominance, employing their baby teeth and the same stereotyped neckbite/shake technique of fighting adults. The one that wins (firstborn has an obvious advantage) keeps the other from nursing until it weakens and dies. Because fighting occurs in the cubs' narrow tunnels, mothers are powerless to intervene. This sibling rivalry kills an estimated 25% of all hyenas in their first month. The surviving male grows faster and is likelier to achieve reproductive dominance; the surviving female eliminates a rival for dominance in her natal clan. There is no reproductive competition between siblings of opposite sex and consequently no killing.

RELATIONS WITH OTHER PREDATORS AND SCAVENGERS

LIONS Very competitive; each tries to take the other's kills. Hyenas usually lose their kills and have to wait for lions to finish eating. However, hyenas in force often unnerve female and immature lions by advancing threateningly and noisily shoulder to shoulder, though rarely attacking. They get nowhere with big male lions.
WILD DOGS Hyenas regularly try to profit from this predator's hunting efficiency, but even when greatly outnumbered, wild dogs usually stand off hyenas by mobbing and punishing one at a time. Lack of cooperative defense places hyenas at disadvantage.
LEOPARD AND CHEETAH Spotted hyenas take their kills at every opportunity.
JACKALS Often annoy hyenas successfully: dart in and steal tidbits from kills; mated couple outwits feeding hyena by moving in from both sides until hyena lunges at one, enabling the other to grab food and run.
VULTURES Hyenas use vulture spotter network to find meat in daytime. When they see vultures volplaning to earth, hungry hyenas race to the spot.

COMMUNICATION

CALLS Among the most vociferous African mammals, with 11 different, intergrading calls.
Groans and soft squeals Often exchanged during greeting.
Whoop Contact call, rising o-o-o.
Fast whoop Rallying call given by excited hyenas (during border conflict or at a kill); brings clan members to the spot. Most whoops are emitted by males and apparently ignored by other clan members. When a female calls, her relatives and offspring react immediately.
Lowing Expresses rising impatience and often leads into fast whooping; voiced by hyenas kept waiting at a kill.

CALLS ASSOCIATED WITH AGGRESSION
Grunting Quiet, very low growl with mouth closed; accompanies aggressive behavior.
Giggling High, cackling laugh, typically emitted by hyena being chased; expresses intense fear or excitement. The call of the "laughing hyena."
Growling Deep, loud rumble, often with staccato vibration: defensive threat voiced by hyena under attack threatening to bite.
Yelling Starts as scream, changing to a roar, voiced by hyena attempting to escape attackers; alternates with growling.
Rattling growl Low-pitched, soft staccato grunts; given as an alarm call-e.g., by hyenas on kill at approach of a lion.

DIFFERENCES IN SPOTTED HYENA BEHAVIOR

Aggression

Parallel walk: two or more hyenas advancing shoulder to shoulder in erect posture, tails brandished over backs; accompanied by lowing and fast whooping. Prelude to mobbing attack on another hyena and attempts to intimidate lions.
Sociable Behavior

Phallic inspection: after mutual sniffing of nose, mouth, head, and neck, 2 hyenas stand head-to-tail and mutually sniff/touch extended phallus for up to 1/2 min., inside leg meanwhile cocked. Part of greeting ceremony between clan members, performed by both sexes beginning first month; but adult males rarely greet with females. Initiative usually taken by lower-ranking individual.

Social grooming: licking and nibble-grooming. Mothers and offspring. Rare between adults.

Hyena whooping. Contact and gathering call.
Courtship

Bowing display: standing behind reclining female with penis extended, hyena abruptly lowers muzzle to ground, advances quickly, bows again, then paws ground close behind her. Retreats immediately if she responds aggressively. Performed by male afraid to approach female he is courting. Pawing deposits scent of the toe gland.

Mating: toward end of 4 to 12 min. copulation, male rests head and body on female.
Mother and Offspring

Weaning tantrum. Thrown by a yearling whose mother refuses suckling.
Response to Competitors/Predators

Mobbing attacks. By spotted hyenas, against rival clans, also against lions, usually in attempt to appropriate or reclaim a kill.

Reprinted from "The Safari Companion" by Richard Estes
nature@nature-wildlife.com