WildlifeCampus – The Behaviour Guide to African Primates 1

Module # 3 – Component # 5

Savanna Baboons
Subspecies
Papio hamadryas hamadryas, Hamadryas baboon
P. h. papio, Guinea baboon
P. h. anubis, olive baboon
P. h. cynocephalus, yellow baboon
P. h. ursinus, chacma baboon

All these subspecies were considered separate species when The Behavior
Guide was published, and still are in some guides, e. g. Kingdon (1997).
Nowadays the consensus among primate systematists is that all the savanna
baboons are a single species and that the hamadryas specific name has priority.
This certainly serves to muddy the behavioural waters, as the hamadryas is the
one subspecies with a very different social/mating system, being one of the very
few mammals with a true harem system (see Primate Introduction, Social
Organization, type number 6) So this account applies to all the savanna baboons
except the hamadryas.

TRAITS

A large, terrestrial monkey with a doglike head (cynocephalus means "dog-

headed").

Weight: up to 50 kg; average of Papio races and species, adult males 23 kg and
adult females 11 - 12 kg, P. h. ursinus from Botswana, adult males 32 kg (27 –
37), adult females 15.4 kg (14 –17).

Length: head and body, ursinus males 72.5 cm (72 - 73), tail 60 - 84 cm;
females 60 cm (52 - 65), tail 56 - 61 cm.

Teeth: razor-sharp 5 cm canines in males only.

Eyes: Small, deep and close-set beneath prominent brow ridge;

Ears: sizeable and nearly naked.

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

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Limbs: sturdy, nearly equal in length, hands and feet short and wide with stubby
digits.

Coat: coarse, hairs brindled, length varying from long, males with well-defined
cape in Guinea and olive baboons, to short in yellow and chacma baboons.

Colouration: reddish brown (papio), olive-brown (anubis), yellow-brown

(cynocephalus), to greenish-brown with blackish lower limbs (ursinus); newborn
black with red face and rump, juveniles distinctly lighter than adults; males darken
and females lighten with age. Bare skin of nose, lips, ears, and extremities black,
eyelids whitish. Callosities and adjacent rump skin same colour as face in adults,
shiny and often with purplish tinge (pinkish in P. h. papio), female sexual swelling
the largest of all monkeys, most extreme in captive animals.

DISTRIBUTION

Papio's adaptation to terrestrial life in grassland habitats has enabled it to become

the most widespread African primate, found virtually throughout the savanna and
arid biomes wherever water and trees or cliffs occur. Forest replacement by man-
made savanna, agricultural expansion, and local extinction of large predators
have enabled the savanna baboon to extend its range.

ECOLOGY

The savanna baboon, in forms distinct enough to be considered different species

by many primatologists, virtually monopolises the broad niche open to a monkey
able to forage equally well in trees and on the ground, whose size and social
organization allow it to venture far from refuges during its daily ranging. The
hamadryas baboon and the gelada are still more terrestrial, subsisting largely on
grasses, but are only able to dominate in, respectively, the most arid, treeless
regions of northeast Africa and Ethiopia's high plateaus. The savanna baboon
dominates everywhere in between there and the rain forest, and in the South
West Arid Zone manages to live in habitats similar to those the hamadryas
occupies in the Somali Arid Zone.

DIET

Savanna baboons utilize virtually all the accessible edible plants within their
range, supplemented by animal foods. Detailed foraging studies in different
populations all agree that the baboon menu features the most nutritious foods
available. Grasses, flowers, fruits, seeds, buds, leaves, shoots, twigs, bark, sap,
roots, tubers, bulbs, aquatic plants, mushrooms, and lichens are all included. An
incomplete list of the plants eaten by baboons near Cape Town included 94
species. In typical acacia savanna habitat, grasses tend to be much the most
important food source, accounting for over ½ a troop's feeding time in the course

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

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of a year, with leaves, stems, flowering heads, and underground portions taken in
different seasons and growth stages. Early in the rains a troop may eat sprouting
grass almost exclusively. Late in the dry season Nairobi Park baboons depend
almost entirely on the still-nutritious rhizomes and corms of dry grasses, for which
their only mammalian competitors are rooting warthogs.

Among the variety of fruit and vegetable foods baboons find in trees, figs and
acacia pods and seeds are particular favourites.

Pronounced size differences between different sex and age classes lead to some
ecological separation within the troop. Young juveniles venture onto branches too
small to support older animals, and older juveniles pluck fruits and flowers from
the outermost twigs of tall forest trees. Females with small infants often sit on the
ground, eat a higher proportion of grass-seed heads than do other classes, and
feed more on low bushes.

Grasshoppers, spiders, scorpions, and other invertebrates may be important

dietary supplements, especially in more arid areas. Baboons living by the sea
forage in tidal pools and caves, where they feed on mussels, limpets, crabs, small
fish, and the like. Vertebrate prey is seldom deliberately sought but is taken
fortuitously, including fish, frogs, lizards, terrapins, the young of ground-nesting
birds, bird and crocodile eggs, small rodents, hares, and the offspring of
antelopes in the concealment stage. Usually only adult males catch prey the size
of hares and fawns, which they rarely share. But cooperative bunting and food-
sharing have developed into a tradition in one long-studied Kenya troop, with
females and juvenile offspring participating (more under Foraging and Predatory
Behaviour). In South Africa, adult males can become major predators on young
sheep and goats.

LIMITING FACTORS

The availability of water and of secure sleeping sites, in the form of tall trees or
cliffs, is of primary importance in baboon distribution. Although baboons can gain
most of the water they need from their food or in the form of dew, and in some
areas do not regularly go to water, probably most troops do drink regularly.
During droughts when surface water is unobtainable, baboons dig in streambeds
to reach subterranean sources.

SOCIAL ORGANIZATION

Terrestrial, non-territorial, multimale troops composed of resident females and

young organized in kinship groups, each associated with specific males.

Baboons live in troops ranging from 8 to over 200 animals but typically numbering
30 - 40, in which adult females outnumber adult males 2 - 3:1 and immature
classes make up half or more of the total. Very small troops may represent
offshoots of large troops. Home ranges vary in size from as small as 400 ha to as
large as 4000 ha, depending on the habitat and without regard to troop size. The

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

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ranges of different troops usually overlap widely. Troops tend to avoid one
another, although different bands may share the same sleeping sites if refuges
are scarce; competition over roosting space, water, or fruiting trees can lead to
inter-troop aggression.

Baboon social organization is highly complex and variable, with a communication

system capable of signalling subtle gradations of emotion and motivation. As
usual, females make up the stable core of the group. They remain their whole
lives in the same troop (or an offshoot, should fission occur), whereas males
emigrate during adolescence and often transfer repeatedly between troops. A
strict and stable rank hierarchy exists between the adult females and their
offspring. Each mother heads up a kinship group within which her offspring are
ranked according to age in descending order, beginning with the youngest. In
social relations between matrilines, the young take the rank of the matriarch; a
lower-ranking adult female has to defer even to the infant of a superior (cf.
vervet). Initially, deference is enforced, if need be, by the active intervention of the
baby's mother or siblings, but by the age of 2½ years a juvenile female's rank is
fixed for life.

Male offspring remain in the kinship groups and subordinate to their mothers until
about 4, when they become adolescent and experience a growth spurt during
which they develop long, sharp canines and become bigger than and dominant
over all females. Females begin reproducing in their fifth year, before they are
quite fully grown, whereas males have to mature for another 3 - 5 years before
they can compete on equal terms with other males for reproductive opportunities.
It is this quest that motivates males to transfer between troops, staying for as little
as 1 month or as long as 10 years.

Male dominance status and reproductive success are of course linked. Females
tend to be receptive to dominant males, especially to those that have recently
joined the troop. Accordingly, low-ranking males would have most to gain by
emigrating. However, dominance alone does not ensure mating rights. Females
prefer to mate with particular males which are regularly associated with them at
other seasons, with whom they are in effect pair-bonded (more under Sexual
Behaviour). Observations of male-female relationships in several different
populations revealed that each adult female had 1 and usually 2 (sometimes 3)
favourites among the adult males; they were most often near her during daily
ranging, roosted in the same sleeping subgroup, and were regular grooming
partners. Proximity was maintained primarily on the initiative of the females,
which preferred to stay within 15 m of a favourite male while foraging and more
than 15 m from other males.

When females came into oestrus, their favourite males were usually their
consorts, on the male's initiative but with the female's cooperation. In an Amboseli
N.P. troop, fewer than half the 10 adult males had special relationships with the
17 adult females. All of them were fully mature and in the top half of the hierarchy.
The first and fifth-ranked males, allied with 6 and 5 females, respectively, were
associated with the most mothers and infants and were the most successful
reproductively.

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

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Godfathers

Males that associate with females play "godfather" roles to their offspring,
beginning in infancy and lasting at least into the second year, when juvenile
baboons become largely independent of their mothers. Almost all friendly
contacts between males and infants, including holding, carrying, grooming, food-
sharing (e.g., meat scraps), and so on, reflect such relationships. When adult
males come to the aid of juveniles or females that are being bullied by other troop
members, it is on behalf of the individuals with whom they are socially bonded.
Not only do young baboons receive increased protection against the dangers of
injury by other baboons and increased access to resources, but godfathers have
even been known to act as foster parents of juveniles whose mothers died.

Apart from the advantages derived by juveniles, low-ranking females may gain
greatly from the presence of male protectors. Mothers of infants up to 5 months
old often suffer considerable interference and distress at the hands of dominant
females that are strongly attracted to babies in the black phase. Most baboons
(especially juveniles and adult females) find such infants attractive, but only
higher-ranking individuals have the nerve to handle infants clinging to the
mother's front, despite her and her baby's distress. Babies may be roughly pulled
and even kidnapped from lowly females. An infant's cries will bring a godfather to
the rescue. But the mere presence of an adult male is usually enough to inhibit
other females from approaching a mother.

In many cases godfathers must be the real fathers of the infants they sponsor.
According to one study, though, males that were probably not the father were as
likely as actual fathers to play godfather roles. The inference is that males also
derive benefits from godfathering infants other than their own. These may include:

a) establishing bonds with a female that will culminate in the fathering of

future offspring;
b) a means for male immigrants to gain acceptance into a new troop more
quickly; and
c) opportunities to use infant, juvenile, and female associates as buffers in
aggressive encounters with male rivals.

In sum, there is increasing evidence that the reproductive success of male

baboons is affected to a large extent by their nonsexual relationships with
females.

Although sexual competition may have been the driving force that caused male
baboons to become twice as big as females and develop lethal weapons, these
same attributes enable troops of baboons to venture away from the safety of
trees and cliffs, despite the presence of large carnivores. Indeed, a leopard is
more than a match for a single male, but the readiness of males to join together in
defending their troop (Altmann and Altmann 1970, Saayman 1971, N. Monfort
pers. comm.) could well account for the fact that even lions seldom molest
baboons (J. Popp pers. comm., author's observations). A group defence
represents cooperation rather than competition, but there is a precedent; males
form coalitions in order to gain an advantage over more dominant males. By

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

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joining together to protect the troop against predators, they are at the same time
safeguarding their own reproductive investment.

ACTIVITY

The baboon is a diurnal monkey which usually becomes active well after dawn
and retires to a secure sleeping place before dark. The troop is a cohesive unit
which keeps together during foraging and the activities of the members are highly
coordinated - 90% do the same thing at the same time. Social (primarily
grooming) and sexual activity are most frequent before and after the day's
foraging. Juveniles remain more or less continually active; their antics start before
and end after all other baboons have begun and ceased stirring.

Unlike most other African wildlife, baboons do not have feeding peaks early and
late in the day divided by a midday inactivity period. Although they may drowse in
the shade when it is very hot, what sets this species apart is the lack of any set
pattern that holds for different troops and populations. A given troop may be
active at any time of day. Baboons have been known to get started before dawn,
to retire half an hour after dark, and even to leave their roosts on moonlight nights
to feed in fruiting trees or to raid crops. Distribution and availability of water, food,
and sleeping sites largely determine how much time a troop has to spend
foraging and travelling in a day, and how much will be left over for resting and
socializing. In Amboseli, cumulative rainfall was found to be a good predictor of
the length of a troop's day journey and of the time the baboons spent feeding.

Day ranging was longest (6 km) in dry months and shortest during the rains (4 -
4.5 km). More than 25% of the day was spent walking and 52% - 55% feeding
during the dry season, compared to 17% - 19% walking and 40% - 42% feeding
in the rains. Daily ranging of 5 - 6 km is about average for baboons living in
acacia savanna; those living in richer habitats such as semi-deciduous woodland
and gallery forest travel only half as far. A troop living in the semi-arid Northern
Province of South Africa made day journeys of 10.5 km. But the record may
belong to hamadryas baboons living in the Ethiopian semidesert lowland: one
troop averaged 13.2 (9.2 - 19.2) km per day.

POSTURES AND LOCOMOTION

Adult males and females move and carry themselves differently enough that a
knowledgeable observer can sex baboons without other clues. More subtle
differences between same-sex baboons reflect an individual's self-confidence and
social rank. However, tail carriage is not a reliable indicator of rank, as sometimes
asserted. It is an indicator of age: the lower part of the tail stands more erect and
the bend becomes more acute with age. The "riding whip position" is diagnostic of
the genus Papio, but there are also differences between species and subspecies
in the amount of bend. The hamadryas baboon and the Guinea baboon carry
their tails in a smooth curve, whereas the tails of other races of savanna baboon
have more kink.

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

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The way a baboon sits depends on how alert it is, ranging from the hunched
squat of a relaxed animal, hands resting loosely on the knees, to the straight back
with head raised and turning of a vigilant baboon (fig. 26.2). Lying positions vary
from flat on the stomach on a branch with arms and legs dangling, to flat on the
back with limbs in the air. The most abandoned poses are seen during grooming
sessions (fig. 27.10).

Baboons move in the usual cross-walk. Running gaits are comparable to the
canter and gallop of a horse, hindlegs and forelegs moving in pairs. Although the
limbs are somewhat modified for terrestrial life, baboons still possess impressive
climbing skill, seeming indeed hardly less competent than forest monkeys after
allowing for the difference in size. Baboons associated with redtail and colobus
monkeys in a Uganda forest could jump as far and were equally agile. However,
the baboons jumped to the ground when seriously alarmed and fleeing, whereas
the monkeys fled through the trees. Equally surefooted on cliffs, baboons ascend
steep places with short leaps and grope for handholds on sheer surfaces. Falls
are very rare, except during play, and baboons land catlike on all fours; one
female dropped 10 m and landed undamaged on a bush-covered ledge. In
Namibia playing juveniles somersault down sand dunes.

Despite their stubby, calloused fingers, baboons have exceptional manipulative

skill. They pluck grass and pick up small objects between thumb and index finger,
and use their fingernails to dig through hard soil and extract roots or beetle
larvae. Actually, the index finger and thumb are the only digits that can be moved
independently, although the fingers are flexible enough to bend back at right
angles to the palm. The hand can be bent under more but the forearm rotates
less than ours.

FORAGING AND PREDATORY BEHAVIOUR

Baboons feed from hand to mouth and often sit while eating. There is no
evidence of tool use; seashore baboons crush shellfish with their teeth rather than
break them open against rocks. Similarly, Serengeti males use their canines to
pry open river oysters imbedded in rocks exposed during extreme low water; the
bivalves burst open with a loud pop (A. Root pers. comm.). But when digging up
grasses to eat the rhizomes, baboons brush or knock off adhering dirt against a
forearm. Invertebrate prey is sought by turning over rocks, logs, and dung, tearing
off bark, groping in holes, and digging up the ground. When eating sizeable
vertebrate prey such as hares or antelope fawns, baboons use their incisors to
strip meat from bone and begin eating at the underside. Such prey is seldom
deliberately hunted, but in a troop where hunting has become a tradition
(discussed under Ecology), males cooperate in stalking and chasing game, and
even share meat.

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

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SOCIAL BEHAVIOUR
COMMUNICATION

Baboon communication is very similar to that of other cercopithecine monkeys.

However, the displays of baboons and the closely related macaques are more
variable than those of other monkeys. They have more expressive faces and
highly graded vocal signals. To understand a given behaviour, it is necessary to
recognize seemingly trivial differences in posture, movements, facial expression,
and accompanying vocalizations, for these register subtle changes in the animal's
emotional state and intentions. To present a list of displays is therefore to make
artificial subdivisions of what is actually a continuum of visual and vocal signals.
The more subtle visual signals are not even listed. For instance, small changes in
normal locomotion (change of pace, halting, looking around) by high-ranking
troop members give the signal for a troop to set off on the day's foraging or to
change direction.

Visual Communication (Agonistic Behaviour)

Dominance/threat displays: staring, with eyebrow-raising and rapid-blinking,

tooth-grinding*, canine display* (tension yawning), penile display*, bristling,
pushups, slapping ground, rubbing ground, branch-shaking, rearing on hindlegs,
lunging, and charging.

The asterisks identify displays performed only or primarily by adult males. Canine
display (fig. 27.11) or threat-yawning is often seen when a lower-ranking male
harasses a superior that is preoccupied with an oestrous female or meat. Unlike
tension or displacement-yawning (which is seen in all classes), threat-yawning is
accompanied by eyebrow-raising, ear-flattening, and other aggressive gestures.
Tooth-grinding can be heard when 2 males are threatening each other at close
quarters. Advance warning of aggressive intent or mood can be read from an
attitude of tension, as when a dominant male assumes an alert posture in a
context other than danger, especially if he stares at someone and slaps the
ground; or if a seated animal gets to his feet and stands with hindlegs planted,
arms rigidly straight, hair on end, and stares with raised muzzle. Adult males
positioned on vantage points and keeping a lookout while the rest of the troop is
foraging often sit with thighs spread and flesh-coloured penis prominently
displayed (fig. 26.2). This behaviour may be a signal to outside males that an
adult male is present and guarding the troop females.

The commonest and mildest form of intimidation is supplanting, when a dominant

baboon approaches and occupies the place of a subordinate.

Submissive displays: avoidance, social presenting, rapid-glancing, fear grimace,

undirected staring with eyes wide, erecting tail, crouching, lying prone.

When a subordinate baboon passes close to a feared dominant, averting the

gaze or undirected staring, the fear grimace, and ear-flattening (as during threat)
are often combined. Grimacing with open mouth, combined with yakking (gecker),
is a more intense expression of fear and submission. A baboon that is cornered

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

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during a chase crouches or lies fiat on the ground, often rigid with fear, while
grimacing and churring.

Social presenting (fig. 26.4) is seen in several different situations, each with some
postural variations.

a) Brief presenting by non-oestrous females and by juveniles, typically to a

dominant male; females approached by males other than regular
companions present and make submissive gestures, which may elicit
genital inspection and mounting by the male.

b) Adult female to a female with a black infant, with hindquarters sometimes

lowered, looking over the shoulder and lip-smacking.

c) Invitation to a juvenile to be carried (fig. 26.4).

d) Lowering the hindquarters by one adult male to another – typically

subordinate – male , which usually grasps the presenter's rump but rarely
mounts. The hindquarters are lowered more in (b) and (d) than during
sexual presenting.

Tension, uncertainty, or displacement activities: yawning, scratching,

twitching (head, arms, shoulders), shoulder-shrugging, muzzle-wiping with hand.

These and other motor patterns are commonly classified as displacement

activities when performed out of context and often in a jerky, desultory way.
Shoulder-shrugging and muzzle-wiping may appear after a baboon has been
startled, as by the sudden appearance of a snake. The male of a consort pair
which is being harassed by rivals may make rapid, sometimes frantic grooming
movements and hurry copulation. Mounting and even the act of copulation may
be performed as a displaced activity by a disturbed male.

Friendly or reassuring displays: grooming solicitation, lip-smacking, greeting,

standing bipedally before another baboon.

Behaviour in this category usually goes together with or leads to body contact
(more under Tactile Communication; see also under Vocal Communication, e.g.,
lip-smacking).

Tactile Communication

See also under Sexual and Parent/Offspring Behaviour: social grooming, nose-to-
nose greeting, social mounting, embracing, touching side.

The greatest part of baboon social interactions is invested in social grooming. The
complicated web of relationships, affiliations, and rank order within a troop can be
deciphered by observing who grooms whom and how (relaxed or tense). The
baboon's day typically begins and ends with long grooming sessions; sleeping
subgroups become grooming subgroups as soon as they come down from their
roosts. Mothers groom their own infants more than they groom any other baboon

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

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and receive more grooming from other females than they give while their infants
retain the natal coat (see under Social Organization). Otherwise, the direction of
grooming is usually from lower to higher rank, males receiving much more
grooming than they give (except to oestrous females) (fig. 27.10). Social
mounting, embracing, and touching the side are frequent responses to social
presenting and probably signal friendly reassurance. They are also seen during
aggressive interactions, between coalition partners, and by individuals attempting
to enlist support (by rapid-glancing, screeching, etc.; see under Fighting).

Vocal Communication

Most of the calls listed in table 27.2 grade into one another, for instance grunting
into barking and roaring, screeching into yakking (repetitive churring).

AGONISTIC BEHAVIOUR

Fighting and Overt Aggression

The frequency and intensity of aggression seen in a baboon troop vary greatly,
largely as a result of troop composition and relationships between individual
males, in particular. If the rank hierarchy is well established, dominance is
asserted and acknowledged with little or no overt aggression. But troops with
recent male immigrants or newly matured resident males tend to be unstable.
Dominant newcomers may be unusually tyrannical and sexually active. Rivalry for
dominance may include frequent threat and counter-threat, the formation of male
coalitions (2 males joining forces to defeat a third, more dominant one) (fig.
27.12), the use of infants to buffer aggression of a rival, and sometimes fighting.
The male's slashing canines make fighting extremely dangerous and
correspondingly infrequent. Yet most adult males bear scars, especially about the
head.

Even in stable troops, practically any disturbance, internal or external, is apt to

provoke disciplinary action by dominant males. Teasing or bullying of small
baboons by older juveniles and squabbles between females frequently trigger
chases punctuated by shrieks, barks, and roars. If a male catches his victim, he
may seize it, beat it with his hands, rub it on the ground, bite it in the neck, and
even pick it up in his jaws. Despite their apparent ferocity and the disparity in size,
such attacks rarely last more than a few seconds or result in visible injury.
Attempts of dominant males to herd the troop away from an external disturbance
(predator, another troop) may result in generalized aggression as the males
canter, double back, and race from side to side to move the females and young
and round up strays.

Just as the incidence of aggression within troops is highly variable, so do

relations between troops vary, apparently depending on the degree of competition
for the same resources (e.g., roosts, fruit trees, water) and the existence or
otherwise of social ties: baboons from troops that have recently fissioned may
mingle and socialize peacefully when they meet. The most spectacular displays

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

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of baboon aggression occur during hostile encounters between troops, when

nearly all the assembled baboons may join in a scene of pure bedlam. Normally
hostilities proceed no further than noisy squabbling, in which males take the lead:
jumping about, branch-shaking, barking, herding their own feisty females and
young away from the skirmish line. But sometimes from 3 to 10 males of one
troop make a mass charge into the ranks of the rival troop, followed by withdrawal
with or without a countercharge by males of the other troop. There is rarely any
physical contact, but the troop with fewer adult males is usually the first to
withdraw.

REPRODUCTION

Females begin menstruating at 4½ -5 years, conceive about a year later, and

thereafter produce young at roughly 1½ - 2 year intervals. Mothers do not
normally resume cycling for 10 - 12 months after giving birth (minimum 5 months)
and typically experience 4 - 5 cycles before conceiving again. Given an average
cycle length of 36 days, a 6-month gestation, and a life span of 20 - 30 years in
the wild, it can be calculated that female baboons spend half their adult lives
taking care of dependent offspring, another third pregnant, and the rest
menstrual-cycling.

Female reproductive condition is plain to see in Papio from the coloration and
degree of swelling of the sexual skin (fig. 26.5). During pro-oestrous (7 days), the
perineum is black with a pink tinge and beginning to inflate. Oestrus follows,
lasting c. 10 days during which the sexual skin is bright pink and fully swollen.
Fading and deflation begin 2 - 3 days after ovulation and end in 7 days with the
sexual skin black and flat, a condition (anoestrus) that lasts for 9 days, when pro-
oestrous begins anew. In the interim, menstruation occurs but seldom with any
external signs of bleeding. In the event of pregnancy the naked skin over the hips
and around the callosities turns from black to scarlet within a few weeks; the
sexual skin is not involved. The sexual skin of lactating females is as in
anoestrous (black and flat). The size of the oestrous swelling is small in
adolescent females and tends to increase with age up to 5½ - 6 years. But the
enormous swellings sometimes seen in captivity, perhaps resulting from repeated
cycling without pregnancy, are not seen in wild baboons.

SEXUAL BEHAVIOUR

Females in pro-oestrous and oestrus can be singled out of a troop of foraging

baboons by behavioural clues even before their posteriors become visible. While
other females remain seated except to move from one spot to the next, an
oestrous female moves about restlessly, and either approaches and solicits
different males or has an adult-male consort. Females in pro-oestrous often
approach and present to several males within a few minutes, standing with tail
bent to one side and looking back over the shoulder, sometimes lip-smacking and
showing the eyelids (fig. 27.13). Females in this stage may allow subadults and
even juveniles to mount them. It is the female, then, which takes the initiative prior

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

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to entering consortship with an adult male, after which either partner may take the
initiative.

Although inflating and swollen females present equally often, males are more
likely to copulate with those in oestrous than in pro-oestrous, and rarely mount
adolescent females (which have small swellings). Even 4 out of 5 presentations
by a tumescent female may only elicit a touch, lip-smacking, or brief grooming, or
no response at all. Males become more discriminating with age; mature
individuals rarely mount and virtually never ejaculate into any but fully swollen
females. Conversely, mating opportunities for subadult and large-juvenile males
are limited to females in the inflating stage. Although a keen interest in a
copulating pair is evinced by the presence of onlookers, there is surprisingly little
evidence of sexual jealousy among adult males.

The existence of pair bonds may be the cause or the effect of this tolerance. In
any case, it has become apparent (from observations by female observers) that
female preference influences the behaviour of consort pairs more than was
previously assumed (from observations by male observers). Females consorting
with highly preferred partners cooperate with efforts to herd them away from other
males, and groom and present to their favourites significantly more often than to
less-preferred partners. For their part, preferred males follow and herd favourite
females more closely, and mount females that strongly prefer them more than
females with no preference for them. In short, the level of a female's receptivity
closely reflects her preference, which in turn affects the male's sexual
performance.

Copulation. The copulatory positions are the same as in other cercopithecine

monkeys, the female standing as during sexual presenting, with tail deflected to
the side (fig. 27.13). However, there appear to be some differences between
subspecies. Chacma baboon copulation consists of a series of mounts, ranging
from 3 in 11 minutes to 9 in 191 minutes, culminating in a mount with an
ejaculatory pause after an average of 10 pelvic thrusts (4 - 20). Females give the
grunting copulatory call after a series of thrusts, then as the male dismounts
typically leap away while continuing to call. Grooming occurs after about half the
copulations. Olive and yellow baboons, by contrast, usually ejaculate during a
single copulation lasting 8 - 20 seconds, after an average of 6 thrusts, with
intervals of at least half an hour between copulations. Females rarely call.

PARENT/OFFSPRING BEHAVIOUR

No strict birth season has been noted, but conception peaks tend to occur during
periods of food abundance (rainy season / summer). Most births occur at night
while baboons are roosting and therefore, although wet infants carried by females
with bloody behinds, hands, and mouths may be seen in early morning,
parturition has rarely been observed in the wild. One female was seen to go into
labour after the rest of her troop had gone to roost. She alternately jerked her
limbs and lay on her side on the ground, but gave birth only after ascending the
trees at dusk. A captive female laboured nearly 7½ hours and delivered in a
squatting attitude while gently pulling on the infant's head. Afterward she licked it

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

WildlifeCampus – The Behaviour Guide to African Primates 13

clean and ate the placenta. It was nursing within 23 minutes of the appearance of
the head.

Selection may favour night births because of the difficulty exhausted mothers
have in keeping up with the troop, especially while supporting a newborn infant
with one arm (clamping its mouth on a nipple gives the baby an additional
anchor). The bright pink skin and black hair of the neonate contrasts strongly with
adult colouration; the male's penis is very conspicuous (adult handlers almost
invariably turn infants upside down and closely examine the genitalia). This
colouration begins to change in the third month and adult pelage is acquired by
the sixth month, although males retain a pink scrotum for at least another year.

From a completely dependent and helpless start, an infant baboon gains

locomotion skill by the end of a month and during the second month develops
enough climbing skill to clamber over small logs and adult baboons. Riding in the
jockey position (fig. 27.14) begins at 6 - 12 weeks, but some baboons never
change from the ventral position. During the second month babies develop the
skill to pick up plants and other objects by hand that they had previously picked
up in their mouths. In months 3 and 4 baboons begin interacting with other infants
that are their companions in the maternity subgroups formed by mothers with
small young. Although still dependent on their mothers for virtually all food and
transportation, they have begun to feed on the more accessible foods and can
climb about within a tree fairly well. Yearling baboons are quite independent and
could theoretically survive if their mothers should die. Yet the youngest surviving
orphans in the Amboseli study (from which the above observations are drawn)
were 17, 20, and 21 months old. Evidently the continuing protection and guidance
of the mother are essential for some time after nutritional independence, despite
the care provided by godfathers.

ANTIPREDATOR BEHAVIOUR

Considering that baboons take extra risks by venturing into open grassland far
from refuges, a correspondingly higher predation rate would be predicted. Yet
nearly all baboonologists have commented on how little predation they have
observed. Some observers maintain that males do not protect females and
young, but only worry about their own skins (R. Harding pers. comm.). If females
and young were really unprotected and vulnerable whenever they left the shelter
of the trees, surely the larger predators would take advantage of the situation?
The fact that they do not is indirect evidence that the presence of adult males
deters attack. There is also direct evidence that not only adult but also subadult
males, and even adult females, will confront predators as large as leopards when
there is no alternative.

The way baboons respond to a predator depends very much on its identity and on
the circumstances. The response of a troop to the presence of a distant lion or
leopard depends on whether they are far from or near a secure refuge. An
attempt to come between a troop and the safety of a grove of trees can create
panic. In contrast, baboons are more relaxed while in short grassland than when

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus

WildlifeCampus – The Behaviour Guide to African Primates 14

moving through cover that could conceal a predator, and are most cautious when
going to water, approaching, a few at a time, only after reconnoitering, while adult
males often stand guard in trees or other vantage points.

When a foraging troop is alerted to danger by an alarm bark (often given by some
juvenile or subadult foraging on the troop periphery), adult males move in the
direction of the disturbance and proceed to demonstrate typical vigilance
behaviour. A sudden disturbance nearby provokes a barrage of loud barking and
a stampede for cover, which would probably confuse a lurking predator's attempt
to single out a victim if its initial rush had failed. The whole troop then makes good
its escape while males and sometimes adult females form a rearguard and, if
necessary, go on the offensive. At least half a dozen cases are known in which
leopards have been mobbed and severely wounded by baboons (N. Monfort,
pers. comm.).

Whereas the leopard is probably correctly regarded as the most effective baboon
predator, lions, spotted and striped hyaenas, chimpanzees, pythons, large
eagles, and humans are also known to have killed and/or fed on baboons. In the
presence of man, baboons rarely put up any defence (but they have been known
to threaten people holding screaming infant baboons – J . Popp pers. comm.).
They run away silently and will jump out of their sleeping trees up until dark. At
night, however, baboons will usually not budge, but manage to discourage
intruding humans very effectively with a rain of stinking excrement (author's
observations).

SOURCES

Altmann 1980, Altmann and Altmann 1970, Bolwig 1959, Bolwig 1963, Busse and
Hamilton 1981, Crook and Aldrich-Blake 1968, DeVore and Hall 1965, Gillman
and Gilbert 1946, Hall 1962a, Hall 1962b, Hall 1966, Hall and DeVore 1965,
Hamilton, Buskirk & Buskirk 1975 , Hamilton, Buskirk & Buskirk 1978, Harding
1976, Harding and Strum 1976, Hausfater 1975, 1977, Kingdon 1971, Kummer
1968, Marler 1965, Napier and Napier 1967, Packer 1979, Ransom 1981,
Ransom and Ransom 1971, Rasmussen, D. R. 1979, Rasmussen, K. L. R. 1983,
Rowell 1966, Rowell 1967, Rowell 1972, Saayman 1971, Smithers 1971, Smuts
1983a, Smuts 1983b, Smuts 1985 , Stein 1984, Stolz and Saayman 1970, Strum
1975, Strum 1981, Strum 1987, Wickler 1967.

Savanna Baboons (Guinea, Olive, Yellow & Chacma) © WildlifeCampus