WildlifeCampus – The Behaviour Guide to African Carnivores 1

Module # 3 – Component # 3

Brown hyaena
Hyaena brunnea

TRAITS

The southern equivalent of the striped hyaena.

Height and weight: 78.7 cm (70.6 - 86.8) and 39 kg (35 - 50), no significant
difference between the sexes. Muzzle broad and short, robust teeth adapted for
cracking bones; long, pointed ears; long legs.

Coat: shaggy with tail and mantle of much longer hair.

Colouration: mostly dark brown, partly obscured by mantle of straw-coloured

hair, legs dark yellow-brown with black stripes; cubs gray, maneless, with more
stripes on body and legs.

Scent glands: large anal pouch that produces 2 secretions (white and black).

Mammae: 4.

DISTRIBUTION

South West Arid Zone and adjacent dry savanna south of the Zambezi River. It
has been exterminated in South Africa except in northern districts of the
Northern and Cape Provinces.

ECOLOGY

The dominant large carnivore in the South West Arid Zone, the brown hyaena
is an opportunistic forager that manages to survive in country where game and
water are both very scarce by eating almost everything with any food value
except herbage: some 58 different kinds of food have been identified in its
droppings. It kills only c. 6% (by weight) of the food it consumes, and is purely a
scavenger of large mammals, the richest but least dependable food.

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Shortages of preferred vertebrate food are made up by eating insects and

various fruits and vegetables. In Namibia the brown hyaena penetrates the
Namib Desert to the Atlantic and scavenges a living on the Skeleton Coast,
where it is called the Strandloper.

Since most of the large mammals that live in the arid zone are migratory, whereas
the brown hyaena is resident, its diet may change radically between the wet
and dry season. In Botswana's Central Kalahari Game Reserve, springboks, red
hartebeests, and wildebeests immigrate during the rains (November – April),
joining the resident gemsboks. Lions and cheetahs and occasional spotted
hyaenas come with them, and the brown hyaena population lives well on the
remains of their kills. In the dry season the proportion of large-mammal remains
in brown hyaena scats falls to only 17%, while the percentage of fruit and
vegetable foods increases, especially tsama and other wild melons, the main
source of moisture during 8 waterless months. Presumably these cucurbid fruits
have nutritional value, too, for brown hyaenas also eat them while water is
available, when they may drink 1 - 3 times a night.

RELATIONS WITH OTHER CARNIVORES

Though it is seldom a predator on big game, the brown hyaena is large and
aggressive enough to rank fourth in the carnivore hierarchy, and effectively
second in the most arid parts of its range, where spotted hyaenas are few and
wild dogs even rarer than usual. Brown hyaenas show no more respect for
leopards than for cheetahs: a lone female robbed a male leopard of the
springbok it had just killed and when the leopard tried to reclaim it, chased it up a
tree. Yet the same species keeps at least 200 m from lions on kills and allows
½ hour after the lions leave before moving in.

SOCIAL ORGANIZATION

Kinship groups including sons and daughters; communal suckling and

provisioning at the den; foraging alone, clan members mark and defend territory
against same-sex neighbours; females mate with nomadic males.

Long considered a solitary species, the brown hyaena has one of the more
advanced and elaborate carnivore social systems, as shown by an 8 - year
study in the Kgalagadi Transfrontier Park, formally the Kalahari Gemsbok N.P.
(KGNP) (Mills 1990) and a 5 - year study in the Central Kalahari Game Reserve
(CKGR) (Owens & Owens 1979). From 4 to 14 hyaenas, including 1 - 4 adult
females, 0 - 3 adult males, 0 - 5 subadults, and up to 4 cubs, live in very
extensive home ranges of 235 - 480 km2 (330 km2, mean of 6 clan ranges) in the
KGNP. In the less arid central Kalahari, the main study clan (average of 13
hyaenas, including 5 females) had a comparatively small range of 102 km2 .
Range size is apparently determined by the distribution of food, whereas
clan size is correlated with food quality and abundance.

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Not only female but also male offspring may remain in the maternal group
after maturing at 2½ years, but whether this is the usual arrangement remains
uncertain. In a sample of 9 male offspring born into 2 KGNP clans, 4 disappeared
as subadults (beginning at 22 months) and 5 remained as adults. One of these
was still living at home 6 years later, but 3 others left within a year. However, only
1 of 9 adult males of known origin was observed to join a group of unrelated
hyaenas. In any case, resident males generally show no sexual interest in
females of their own or even other groups, and show little antagonism toward
nomadic males. These, representing 33% of the adult males and about 8% of the
total population, wander through the clan territories and mate with any oestrus
females they encounter.

But both resident males and females behave intolerantly toward trespassers of
the same sex from neighbouring territories, often chasing and neck-biting those
they catch (see Fighting). Clan members post their territory and at the same time
communicate with one another by scent-marking throughout their range (see
Olfactory Communication).

Apart from the submissive behaviour shown by the young toward adults, no
evidence of a rank hierarchy was noted in the KGNP study, whereas a definite
rank hierarchy was described in the CKGR study group, which was reinforced at
every meeting through ritualised displays of aggression and submission. A
male was the alpha animal but otherwise dominance was not sex-linked. The
presence in the group of 3 unrelated immigrants, 2 females and a male, could
have caused intra-group relations to be unusually competitive. The greater
seasonal abundance of medium and large carcasses, combined with a smaller
range, would also increase food competition and frequency of meetings
between individuals. Two or more hyaenas were present at over half of 87
carcasses of springbok or larger size, and occasionally up to 6 gathered and
socialized between bouts of leisurely feeding. Even adults that were unrelated
and unmated often (at 23 carcasses) fed together; this would seem to reflect the
same tolerance of nomads and individuals of opposite sex shown by resident
KGNP hyaenas. However, no more than 3 hyaenas fed together at the same
time; the rest had to wait, sometimes in vain.

Communal Suckling and Provisioning of Offspring

Several months after whelping, clan females bring their young to a centrally
located den where the cubs are reared communally. Four females in the
CKGR clan had 5 cubs between them, ranging in age from 4 to 20 months. They
all suckled one another's young and brought food back to the den (a springhare,
birds, a jackal carcass, carrion, tsama melons, etc.). Even a cubless female
brought food and tried to suckle cubs. Related subadults of both sexes also
brought food, but sometimes scavenged others' offerings themselves. Females
and subadults would often spend several hours a night sleeping and socializing
around the den, but no adult remained on guard; they spent the days in favourite
resting spots up to several hundred meters distant. (More under Parent/Offspring

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Behaviour.) The adult males did not provision the young and none was seen
at the dens.

In the southern Kalahari population, there was rarely more than 1 litter of cubs in
a den at a time and even when 2 females had offspring they only occasionally
suckled each other's cubs. Adult males helped provision cubs to which they
were related.

ACTIVITY

In KGNP, brown hyaenas were active 80% of the time from 18h00 to 06h00 and,
walking at the rate of 4 kph, travelled a distance of 31 km (1.2 - 54.5) a night in
search of mostly small food items. They invariably foraged alone and found
vertebrate food items or the nutritional equivalent (10 wild fruits) every 9.2 km.
Resident spotted hyaenas, in contrast, foraged mostly in groups averaging 3
animals and had to move a mean distance of 32.7 km to make a kill or find a
nearly intact carcass. Brown hyaenas of the CKGR travelled as far as the more
southerly population in the dry season (20 - 30 km a night), but only 10 - 20 km
during the rains, when they had 2 activity peaks from c. 19h30 to midnight and
02h30 to dawn, with an intervening rest period.

FORAGING BEHAVIOUR

A foraging individual quarters the ground in a zigzag course with head raised
and ears erect, relying mainly on its sense of smell, especially in dense bush
and tall grass. Often a hyaena will stop and turn abruptly into the wind, head and
muzzle raised and turning from side to side. A female winded a calving oryx 0.6
km upwind, ran to the spot through thick bush, and tried to approach, but desisted
after the oryx's third charge. Hearing - and comprehension - are also
impressive: one foraging hyaena turned and loped 400 m downwind and
appropriated a kori bustard that it had apparently heard 2 jackals kill. Within 15
minutes 2 other hyaenas arrived.

Although brown hyaenas sometimes stalk a bird or hare, they usually make no
effort to move quietly and will pursue small game disturbed by their passage for
only a short distance. Occasionally one will undertake long, zigzag chases after
springhares, which cannot hop very fast but dodge superbly; sometimes a hyaena
manages to dig one from its burrow. But only 9.6% to 14% of observed hunts
were successful.

Brown hyaena © WildlifeCampus

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Food Caching

Brown hyaenas cache surplus food. Some 70% of the times hyaenas were
seen at a large carcass, individuals carried a piece, preferably a hindleg, 100 -
600 m and cached it in a thicket, usually returning to eat it early the next night.
One animal may remove up to 3 legs before any competitors discover the kill.
Other hyaenas and jackals seldom find a cache, even though the owner scent-
marks bushes and grass within 15 - 20 m of the site. Hyaenas may also carry
scavenged items, mostly small game, back to the communal den instead of
caching them.

In the KGNP study, 26 out of 40 times that a hyaena was seen to find suitable
food, all or part of it was carried back to the den, an average distance of 6.4 km,
and equally often by males and females.

SOCIAL BEHAVIOUR COMMUNICATION

Olfactory Communication:

Scent-marking with anal-sac secretions and dung; presenting anal gland during
greeting ceremony (described under Agonistic Behaviour).

Foraging brown hyaenas pause 2 - 3 times/km to paste grass stems, bushes,

and other objects with anal-sac secretion. The locations of scent-marks 3
adult males and 2 adult females were seen to deposit within their territory in 1
year are shown in figure 20.8. This clan left an estimated 145,000 marks
throughout their property, of which at least 20,000 would be potent at any given
time (the odour being detectable to the human nose for 30 days minimum).
The highest concentration of marks was in the central, most-used part of the
range, especially along trails and near water holes, dens, latrines, large
carcasses, food caches, and other focal points. Marking density decreased in the
outer part of the range, although the rate of pasting increased when hyaenas
visited the borders, as it did during and after social interactions. Through
computer modelling, it could be shown that intruders into a territory would be
likely to encounter an active scent-mark within the first 250 m.

In addition, brown hyaenas maintain latrines where up to 15 defecations may

accumulate. The distribution of some 85 latrines that were found in the above
territory was more clumped than random, and groupings of latrines occurred
mainly near the border. The majority were situated next to a tree or bush (75%),
or other landmark such as the side of a road (15 %) or pool (2%). Furthermore,
24 out of 34 randomly chosen latrines beside Boscia trees were situated on the
south side. Thus latrines were specifically associated with landmarks and
those on the border were visited more frequently by clan members than interior
latrines.

Brown hyaena © WildlifeCampus

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Hyaena brunnea deposits 2 different anal-sac secretions in turn, first the usual
white pomade, then a black secretion (see fig. 20.2 and description of pasting in
striped hyaena account). The white secretion is produced by sebaceous
glands and its odour is long-lasting; the rather watery dark secretion is
produced by apocrine glands and its smell fades faster. Gas-liquid
chromatographs of the secretions from different hyaenas show consistent
differences in the concentrations of the constituents, which could serve for
individual identification. There is also behavioural evidence that brown hyaenas
can distinguish between the marks of clan members and outsiders. The
secretions probably function not only to establish territorial rights but also as
chemical messages among pack members which reveal both who made the mark
and how long ago. One important benefit may be to avoid foraging in areas
already covered by other hyaenas.

Tactile Communication

Members of the same social group occasionally lick or nibble one another,
concentrating on the neck region. Performed most often between subadults and
adults, mutual grooming probably has a social as well as a sanitary function
(reaching an inaccessible spot), serving to reinforce social and kinship bonds.

Vocal Communication

Squeal, squeak, scream, yell, and growl or grunt; 2 whines, 4 growls, yell, and
hoot.

a. Squeal: a shrill, sharp cry emitted by juveniles or other subordinates while

approaching to greet or beg food of a dominant hyaena.

b. Squeak: a hoarse or rasping cry associated with carpal-crawling (abject

submission).

c. Scream: high-pitched, cackling shriek, given by a hyaena whose neck is being

bitten.

d. Yell: a loud, abrupt, high-pitched call associated with defensive threat.

e. Growl or grunt: low-pitched, breathless, throaty sound given while muzzle-

wrestling. All these calls except the yell are short-range signals audible at no
more than a few hundred meters.

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AGONISTIC BEHAVIOUR

Dominance/Threat Displays:

Bristling, erect posture with ears pointed, scratching ground with forefeet,
growl/grunt.

Submissive Displays:

Fear grimace (fig. 20.4), ears flattened, presenting anal gland. carpal-crawling,
lying prostrate, plus vocal accompaniment (above).

Fighting:

Muzzle-wrestling, neck-biting.

When 2 hyaenas meet, they may either perform the greeting ceremony, ignore or
actively avoid each other, or behave aggressively and sometimes fight.

Greeting between equals begins with mutual sniffing of head, neck, flanks, and
anus as 2 hyaenas stand in reverse parallel, with tails over their backs. An inferior
individual presents the anal gland: cringing, ears flattened, grinning, cackling, it
cuts in front of the other and passes its everted anal pouch under the other's
nose, often repeatedly. Greetings are exchanged mainly between group
members and presenting is most commonly seen when cubs and subadults greet
adults; adults were not seen to present to younger hyaenas.

Behaving submissively has some tangible benefits. A hyaena feeding at a kill

may tolerate another that carpal -crawls up and lies low while eating. And a
hyaena carrying food can usually be forced to surrender it to another that runs up
with ears flattened, body lowered, tail raised or outstretched, and squeals while
latching onto the meat. This behaviour derives from and is mostly seen in
juveniles begging from adults bringing food to the den.

Encounters between members of neighbouring clans of the same sex were

almost always hostile and accounted for all but 2 of the fights observed in
KGNP. However, only 16 fights were seen in 2500 hours of hyaena-watching, and
these involved highly ritualised neck-biting bouts lasting from a few seconds to a
few minutes. They occurred mainly in border areas and the behaviour of the
contestants foretold the outcome from the start: one was always the aggressor,
which bit, pulled, and shook the other by the neck; the other yelled and growled
but failed to bite back. The defeated animal always left the area while the
winner usually remained. Several far more severe bouts were observed in the
CKGR: one between 2 unrelated females that lasted 2¼ hours, during which the
inferior's neck was continuously gripped for up to 18 minutes at a time, and an
engagement of over 3 hours between 2 unrelated males.

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Members of the same group often engage in muzzle-wrestling, which may

represent the contest version of the one-sided neck-biting encounters between
residents and intruders. Standing tall with hackles raised and facing at about a
135-degree angle, 2 contestants throw their heads wildly while biting and pushing
at each other's muzzle and neck, with mouths wide open and teeth showing,
meanwhile growling or grunting. Most bouts involve young animals and
appear playful, especially those between younger and older animals. Females
may dominate males and vice versa, but as usual greater ferocity is shown
toward members of the same sex. Subadults try, often successfully, to engage
adults, which rarely jaw-wrestle with each other. Engagements between adults
and subadults are sometimes serious, the adult becoming increasingly
aggressive while the subadult behaves submissively and whines pitifully.
Subadults of both sexes which were treated in this way subsequently emigrated.

Thus, dispersal of 4 male and 2 female offspring of the CKGR study clan was
preceded by progressively more severe conflicts with same sex adults over
a 3 - 7 month period beginning at c. 22 months.

REPRODUCTION

Breeding is unsynchronised and perennial. Litter size is 1 - 4. The modal number

in 15 KGNP litters was 3.

SEXUAL BEHAVIOUR

Nearly ½ (46%) of observed encounters between clan females and outside males
included sexual behaviour, compared to only 6% of encounters with group-
living males (sniffing vulva or urine, only). Two females were known to mate with
at least 4 different nomadic males during an oestrous period of several days.
Mating is conventional and doglike, featuring some chasing and defensive
aggression, mutual anogenital sniffing, and close-following. During copulation,
which may last 2½ - 10 minutes, the female stands with head low, back slightly
arched, tail raised and deflected. The male rests his whole body on her (see
spotted hyaena, fig. 20.14) and may lightly nip and pull her nape hair. A captive
pair copulated at the rate of 15 times in 9 hours (maximum 5 times in 1 hour)
during the second part of a 2-week oestrous period. In the wild, too, copulations
with different males were seen 9 days apart, between which the female was often
alone.

A combination of factors is thought to account for the strategy of mating

with nomadic males: erratic, often long intervals between oestrus cycles (12 -
23, up to 41 months between litters), very large territories, and solitary foraging all
select against attempts by resident males to monopolize mating with a group of
females.

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PARENT/OFFSPRING BEHAVIOUR

Communal dens may remain in continuous use for several years. In the
CKGR, dens are typically situated on bush-covered dunes preferably
overlooking a river bed. The one occupied by the study clan had 2 entrances
183 cm apart, which narrowed from 152 cm diameter to 50 and 80 cm in the first
½ m, then to 30 - 35 cm. Adults entered the vestibule to rest a few hours or leave
food, but only younger cubs could use the small tunnels, excavated by
themselves. Around the entrances discarded soil made an earthwork where cubs
and adult females spent a lot of time resting while nursing and eating,
monitoring their surroundings, and socializing.

Brown hyaenas give birth in a separate den, transferring their cubs to the
communal den when they are c. 3 months old. Cubs begin to eat meat at 3
months but continue to nurse until at least 10 months old and may only be fully
weaned at 15 months. They suck for 25 - 30 minutes at a time, apparently on a
first-come, first-served basis.

The mother of a 4-month juvenile suckled a 10-month youngster without her own
offspring even being present. Females with small cubs spend up to 5 hours a
night attending them, often paying 2 visits. But after parking them in the
communal den, mothers pay only 1 visit a night and may leave juveniles older
than 8 months unattended for 2 - 3 nights. Thanks to the communal nursing
and provisioning arrangement, these youngsters need not fast while
waiting (cf. spotted hyaena).

With tunnels for safety, the presence of older along with younger cubs
makes adult baby-sitter/guards unnecessary. The smallest cubs make
running dives into the den at the slightest disturbance. Cubs that have
outgrown the tunnels put on a defensive threat display (see striped hyaena
account, fig. 20.5). Those over 14 months old may sleep in or away from the den,
returning each evening. Though they go off foraging beginning at 14 months, the
den remains a focal point through the second year.

PLAY

Cubs of different ages lie in contact, social-groom, and play together. They
hold free-for-alls in which older cubs treat younger ones so roughly that all cubs
acquire neck scars during jaw-wrestling and neck-biting play fights. Yet the
younger juveniles often ask for it by biting the ears, legs, or tails of resting older
cubs, by dancing to and fro before them or flopping onto their heads. By the time
they become adults, brown hyaenas are played out.

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SOURCES

Gorman and Mills 1984. ; Mills 1978. ; Mills 1982a. ; Mills 1982b. ; Mills 1983. ;
Mills 1984. ; Mills 1989. ; Mills 1990. ; Mills, Gorman & Mills 1980. ; Mills and Mills
1978. ; Owens, D. D., and Owens M. J. 1979. ; Owens, M. J., and Owens D. D.
1978. ; Owens, M. J., and Owens D. D.1984. ; Yost 1980.

Brown hyaena © WildlifeCampus