CONTEXTS AND PATTERNS OF INJURIES IN FREE-RANGING

MALE BABOONS (PAPIO CYNOCEPHALUS)

by

CARLOS DREWS 1 ,2)

(Department of Zoology, University of Cambridge, Cambridge, UK)

(Ace. 2-XI-1995)

Summary
Injury in male baboons (Papio cynocephalus) was investigated as an indicator of damaging
fights in order to provide a framework for analyses of conflict resolution and dynamics
of agonistic competition in primates. The vast majority of wounds were canine slashes
resulting from intraspecific face-to-face combat. Wounds were more common in males
than females. In males they concentrated on the right side of anterior parts of the body,
principally the head. Wounds took on average three weeks to heal. Aggressive conflicts
represented 10% of all interactions between males. Less than 1% of aggressive contests
led to injury. The individual rate of injury from fights with other males was on average
once every 1.5 months. The winner of damaging fights was sometimes the wounded indi­
vidual. The number of wounds per damaging fight was not related in a simple way to the
presence of proceptive females or to recent immigration events. Four fights yielding the

1) Present address: Programa Regional en Manejo de Vida Silvestre, Universidad Nacional,

Apdo. 1350 - 3000 Heredia, Costa Rica, e-mail: cdrews@irazu.una.ac.cr
2) I am thankful to the Government of Tanzania, Tanzanian Commission for Science and
Technology, Tanzanian National Parks and Serengeti Wildlife Research Institute for permis­
sion to study baboons in Mikumi and Gombe National Park. The Mikumi Baboon Project,
co-directed by R. Rhine, S. Wasser, and G. Norton, and the Gombe Baboon Project, coor­
dinated by A. Collins and J. Goodall, facilitated the study of habituated baboons and hosted
the author at the respective study sites. The field assistance of A. Sindimwo in Gombe and
of C. Kidung'ho, W. Marwa and A. Njalale in Mikumi was invaluable. I am most grateful
to A. Schmidl-Drews, who collected half of the data for this study. Previous versions of
this manuscript benefited greatly from comments by K. Eltringham, P.C. Lee, G. Norton,
D. Hawkins, A. Whiten, G. Cowlishaw and an anonymous referee. This study was funded
by Kings College Cambridge, Bedford Fund, Durham Fund, Bartle Frere Exhibition Fund,
Gordon Wigan Income Fund, Mary Euphrasia Mosley Fund, Cambridge Philosophical So­
ciety, Sigma Xi, Leakey Foundation, and the Harry Frank Guggenheim Foundation. The
preparation of this manuscript was funded by the Max-Planck Gesellschaft.

© E. J. Brill, Leiden, 1996 Behaviour 133, 443-474

444 CARLOS DREWS

highest number of injuries, however, involved recent immigrations or attempts to immigrate

by adult males in their prime. Contexts of male injury observed during infliction include
challenges to the resident alpha male by newcomers, intertroop encounter, fights over pro­
ceptive females or unusual foods, redirected aggression, defense of a female and a fight
unrelated to any obvious resource. This study and anecdotal reports from the literature point
at various implications of injury to male baboons, including physical impairments which
can constrain feeding efficiency, limit access to resting sites and safe retreats, cause a drop
in dominance rank, jeopardize mating success and even result in death. Severely injured
males typically reduce interaction rates, retreat to the periphery of the troop or emigrate
temporarily. Although most wounds are small and heal well, the potentially high costs
of injury probably exercise strong selection pressure on contestants for means of peaceful
conflict resolution, given that during fights both baboons risk injury irrespective of their
competitive abilities. The potential fitness consequences of inflicted injury can explain the
evolution of the formidable canine weaponry of some male primates.

Keywords: aggression, sexual selection, trauma, competition, mortality, laterality, primates.

Introduction

Most studies on animal fighting have focused on the determinants of out­

come and on the pattern of information exchange during the interaction (see
Huntingford & Turner, 1987). Surprisingly little is known about the con­
sequences of wounds resulting from intraspecific fight, given that the risk
of injury and its associated costs are a central component in game-theory
analyses of animal conflict (e.g. Maynard-Smith & Price, 1973; Maynard­
Smith, 1974; Parker, 1974; Huntingford & Turner, 1987, p. 277 if; Enquist
& Leimar, 1990). There is evidence of death resulting from escalated,
intra-specific fights in a variety of taxa including molluscs, crustaceans,
insects, spiders, amphibians, birds and mammals (examples in Huntingford
& Turner, 1987, Table 3.1 therein). In general, therefore, injuries are as­
sumed to be costly in view of their potentially lethal consequences. Little is
known about the non-lethal implications of injury sustained in fight, how­
ever. In red deer Cervus elaphus stags, recovery from injury is associated
with loss of dominance and antler mass in the subsequent rutting season
(Vogt, 1948), and can lead to a considerable reduction in reproductive suc­
cess (Clutton-Brock et al., 1982, p. 134). There are few accounts from
field studies concerning the pattern, context and implications of lesions.
During escalation animals risk to suffer a potential fitness reduction as
a consequence of injury. There can be direct costs of injury, such as
 INJURIES IN MALE BABOONS 445

e.g. a greater energetic expenditure from limping or the mating opportuni­

ties missed due to a reduction in competitive ability. In addition there are
stochastic costs which mayor may not be incurred, and arise e.g. from an
increased risk of falling prey to predators in the case of a locomotory hand­
icap. In principle, however, damaging fights, defined as contests leading to
injury to any or both participants, need not necessarilly reduce the fitness
of the victim because injury can be minor and may heal rapidly. Thus,
the type of injury, its severity and its healing time largely determine the
kind and magnitude of the handicaps attached to it. The aims of this report
are (1) to present field data on lesions in male savannah baboons (Papio
cynocephalus), (2) to bring together the anecdotal information about male
injuries scattered in the vast literature about baboon behaviour and ecology,
and (3) to evaluate the implications of injury to the victim.
The theory of animal contests has been applied to agonistic competition
in primates (e.g. Popp & DeVore, 1979), but studies of fights are missing in
this order, which thus lacks the empirical data required in some models of
aggressive competition. An insight into pattern and context of damaging
fights can be gained from analyses of injuries, in a similar way to the
use of scars as indicators of aggression levels in rodent populations (e.g.
Christian, 1971; Turner & Iverson, 1973; Rose & Gaines, 1976; Rose,
1979). Primatologists report on the incidence of wounds and at the same
time on the lack of direct observations of wound inflictions (e.g. Boelkins
& Wilson, 1972; Symons, 1978, p. 168). The field worker is more likely
to observe the result of aggression, i.e. injuries, than the aggression itself,
because fights resulting in injury are brief and relatively rare, whereas
injuries can be still recognized several days after the incident.
It is necessary to know how animals fight and the nature of their weap­
onry in order to assess the risk of injury to the combatants (see Geist,
1966, 1978; Packer, 1983; Lincoln, 1994). Fights have been described, for
example, in crustaceans (e.g. Hyatt & Salmon, 1978; Dingle, 1983), insects
(e.g. Davies, 1978; Parker & Thompson, 1980; Thornhill, 1984; Marden
& Waage, 1990), fish (e.g. Laudien, 1965; Simpson, 1968; Dow et al.,
1976; Beaugrand et al., 1991), birds (e.g. Smith & Hosking, 1955; Stout,
1975), pinnipeds (Le Boeuf, 1971; Haley, 1994), ungulates (e.g. Geist,
1971; Walther, 1974; Wilkinson & Shank, 1977; Clutton-Brock et al., 1979;
Fryxell, 1987), carnivores (e.g. Poole, 1973; Havkin & Fentress, 1985), and
446 CARLOS DREWS

mammals in general (Harrison Matthews, 1964). Little has been written

about fights in primates. Two studies have specifically addressed fights
and injuries in baboons. Bramblett (1967) did an osteological analysis of
baboon injuries and diseases. Owens (1975) compared aggression during
play with other aggressive episodes among juvenile and adult olive baboons
(P. c. anubis), and included information on fight and wounding patterns in
adult males. The most common movement pattern in aggressive episodes
between adult males is chasing, lasting on average 12.5 s, about three times
longer than chases between females or between juveniles. The second most
common behaviour is sparring, i.e. partners stand facing each other, grapple
with the arms and hit one another about the head. Less common behaviours
include, mutual and non-mutual biting, non-mutual mauling and wrestling.
Several field studies of baboons make marginal reference to fights and
injuries. This information is cited in the relevant sections below. Analyses
in this report are based on wound records and some direct observations of
damaging fights.

Canines: the baboon s weaponry

During contact fights, male baboons utilize their canines as the main weap­
on to inflict wounds to the opponent. The remarkable degree of sexual
dimorphism in canine length in the baboon is typical of its multi-male
social organization (Harvey et al., 1978; but see Martin et al., 1994). The
exposed shaft of a male's upper canine can measure 2.3 em (Ransom,
1981). The anterior edge is rounded, while the posterior edge and the tip
are very sharp (pers. obs.). Males are believed to sharpen their canines
prior to combat by grinding their teeth (Brain, 1992). The canines are
effective weapons in antipredatory behaviour, for male baboons are known
to kill dogs (Canis familiarisi and attack leopards (Panthera pardus) (see
review of baboon responses to predators in Altmann & Altmann, 1970,
p. 170 fj). In addition, male canines are responsible for injuries on females
(Smuts, 1985, 1987; pers. obs.) and juveniles (e.g. Altmann & Altmann,
1970, p. 46; pers. obs.). Injuries to juveniles inflicted by males can be lethal
(Altmann, 1980, p. 203; pers. obs.). Broken canines are not uncommon and
are probably a consequence of clashes between the teeth of both contestants
(Paterson, 1973; Packer, 1979b; pers. obs.). It is conceivable that the large
upper canines of males have structural adaptations to the lateral impacts
 INJURIES IN MALE BABOONS 447

suffered when parrying the opponent's blows (see description of fights

below). Such adaptations are found in the horns of male antelopes (Packer,
1983). The nails of baboons are small and assumed to playa minor role
in the causation of wounds, with the exception perhaps of injuries to the
sensitive skin of the swollen perineum of proceptive females.

Methods

Four well habituated troops of yellow baboons (P. c. cynocephalus) were observed in
Mikumi National Park, Tanzania. These included the two Viramba troops which have
been monitored in Mikumi since 1975 (see Rhine, 1986). The number of males in a troop
varied from two to five during the study. Baboons in Mikumi were observed on 164 days
in two periods spanning a total of 16 months, between May 1990 and November 1991.
The habitat in Mikumi National Park is characterized by Miombo woodland on the hills,
and open woodland and grassland in the valley. For details of the Mikumi habitat see
Norton et al. (1987). In September and October 1991, one additional troop of habituated
olive baboons (P. c. anubis) with seven adult males, the Beach Troop, was observed on 33
days spanning six weeks in Gombe Stream National Park, Tanzania. Common vegetation
zones in the range of Beach Troop was the evergreen and the deciduous forest, as well as
the beach on the shore of lake Tanganyika. For details of the Gombe habitat see Ransom
(1981), Goodall (1986) and Collins & McGrew (1988). The data set collected corresponds
to 1413 h of baboon observations. The methods and subjects of this study are described in
more detail in Drews (l993b).
All adult and subadult males of the study troops were examined by two observers
(c. Drews and A. Schmidl-Drews) from 2-5 m distance on each observation day for recent
lesions. Scars, mutilations and other permanent marks resulting from wounds past are not
included in the sample of this study. Adult males are defined as fully grown immigrant
males. Subadult males are males over six years, residing in their natal troop and not show­
ing consort behaviour. A total of 123 fresh lesions were recorded on 19 males belonging to
study troops and several other males not individually identified. An additional four injuries
on males were recorded in Gombe. Data for the Gombe olive baboons are only included
in analyses where explicitly so stated. Lesions on adult and subadult females, as well as
juveniles were also recorded.
Each record included the bodily location of the wound, its shape (cut, tear, puncture,
fracture, other), size, age in days or an estimate thereof if knowingly less than eight days
old, healing stage, identity of the individual and the context of the injury if observed. Two
size categories were used in the analyses, Small (i.e. less than 5 em) and Large (i.e. 5 ern or
larger). These categories applied to cuts and tears; the depth of wounds was not recorded.
A wound was considered as healed when it was closed, dry, free of flies and not groomed
any longer. Fractures and other internal wounds on limbs were considered healed when
limping ceased. Fight is used here to describe the section of an agonistic contest, in which
aggressive, bodily contact occurs.
The troop is defined spatially as the most coherent set of baboons containing 90% of
troop members. The remaining 10% are considered with the troop if they are within one
448 CARLOS DREWS

troop diameter of the troop's periphery. Baboons were recorded as peripheral when they
occupied corners of the largest imaginary polygon around the troop. Baboons located more
than one troop diameter away from the periphery of the troop were considered outside the
troop and not included in the polygon.
Definitions of dominance status and rank follow Drews (1993a). Statistical tests are
two-tailed, unless otherwise stated. Chi-square values were corrected for continuity (Yate's
correction) in analyses with one degree of freedom.

Results
Description offights
The following description of fighting techniques is the result of direct ob­
servations of 91 instances of escalated aggression between male yellow
baboons and frame by frame analyses of two video records. In fights,
males use their canines in a slashing rather than biting motion. This con­
trasts with fights among females and juveniles were biting and holding the
opponent with closed yaws was common. Males used their muzzle both
to strike at the opponent and to fend off the opponent's blows. Contes­
tants remained quadrupedal during fights, but used single hands to slap
the opponent, sometimes sweeping its forelimbs away thus inducing a loss
of balance. Exchanges of canine blows and hand slaps, which can lead
to injury to one or both contestants, generally did not last more than 5 s
(also Owens, 1975; Brain, 1992). Complete fighting sequences, which
may include brief grappling bouts, did not exceed 30 s. Although a male
olive baboon was reported to have ridden on its fleeing opponent while
directing bites to its neck region (Popp, 1978, p. 99), male fights in this
study typically ended in either a chase or one male leaving the fighting
ground. Superficially, male baboon fights resemble carnivore fights in that
canines are also used as both an offensive and defensive structure and in
that sweeping in a judo-like fashion is a component of the interaction (e.g.
wolf pups Canis lupus: Havkin & Fentress, 1985).

Types and causes of injuries

The sample sizes differ between analyses below, due to some incomplete
records during the initial stages of the study. Male injuries included cuts,
tears, punctures, fractures and a swollen shoulder. Limping with no external
sign of the cause represented 15% of 104 injuries. The majority of wounds
 INJURIES IN MALE BABOONS 449

TABLE 1. Types of injuries among adult and subadult male baboons

Type of injury Cases %

Cut 67 76
Tear*) 6 7
Puncture 8 9
Fracture 6 7
Swollen shoulder 1 1

Total 88 100%

*) In tears, as opposed to cuts, a piece of skin hangs from the wound exposing it.

TABLE 2. Causes of injury among adult and subadult male yellow baboons

Cause of injury" Cases %

Aggression confirmed 13 13
Aggression suspected 86 8S
Predator attack suspected 1 1
Hit by vehicle confirmed 1 1

Total 101 100%

*) 'Aggression confirmed' is based on the appearance of wounds immediately after observed

fights (typically cuts, tears and punctures). 'Aggression suspected' is based on the type of
injury as known from observed aggression. 'Predator attack suspected' is an open wound
in which a portion of muscle is missing, similar to wounds observed in ungulates during
the study.

were cuts, tears and punctures (Table 1). In complete records, nearly all
wounds (i.e. 98%) could be confidently attributed to canine action during
intraspecific fights (Table 2). The sample of injuries attributed to agonistic
encounters between males is composed of 92 injuries on 29 adults and
seven on two subadults. The cause of injury was unknown in 22 cases (not
computed in Table 2), twelve of which corresponded to limping. Where
size was indicated (N = 82), large wounds made up 27% of the sample. In
the main study troop (Viramba 2), adult and subadult males did not differ
significantly from adult and subadult females in the proportion of large to
small wounds (X 2 = 1.89, df = 1, p = 0.168).
450 CARLOS DREWS

Contexts of injury
The infliction of fifteen injuries, i.e. 12% of all lesions recorded in males,
was witnessed during ten damaging encounters. Eight immediate contexts
leading to injury were identified. Injury occurred (1) in two out of eight
coalition attacks during competition over a proceptive female, (2) in one
out of five dyadic fights over proceptive females, (3) once in a fight during
an intertroop encounter, (4) once in a fight between an alpha male and a
non-resident male attempting immigration (see also Results: Number of
wounds per damaging fight), (5) twice in fights over unusual foods (i.e.
banana offered by passengers from a vehicle and kitchen refuse at the pit
of the tourist lodge, respectively), (6) once in a fight over no apparent
immediate resource, (7) once when a male redirected an attack received
by a consorting male against another male (olive baboons), and (8) once
when an old male, whose upper canines were missing, was wounded as
he defended a non-proceptive female from an attacking, higher ranking
male (olive baboons). Injuries have sometimes preceded the emigration of
male baboons (e.g. Harding, 1980). In this study, the disappearance of a
newcomer male, who had recently taken over alpha status in the troop, was
noticed on a day when two of the three resident males had several fresh
slash wounds.
Injury was neither observed resulting from any of 513 socio-sexual in­
teractions between male baboons nor from any of 345 supplantations over
natural foods or resting places such as shade, suitable branches, stones, or
bare area of ground. Thus, common resources of small relative value were
usually not involved in contests resulting in injury.
All witnessed inflictions of injuries on males were sustained during fights
with other males. On two occasions the winner of the fight injured the
loser. Conversely, in three cases the winner (i.e. the animal which obtained
the contested resource) sustained a small injury while the loser remained
unharmed. During an immigration attempt both contestants received large
wounds. Thus, being the superior combatant in a fight is no guarantee
against injury.

Frequency of injury

Demographic composition data from the main study troop was used to
analyse the distribution of injuries among different sex and age classes
 INJURIES IN MALE BABOONS 451

TABLE 3. Proportional representation of injuries among sex/age classes in

troop Viramba 2

Sex/age class Injuries observed Injuries expected Mean daily % in troop"!

Adult males 35 (45%) 8.7 ILl %
Subad. males 7 (9%) 4.6 6.0%
Ad. + Subad. males 42 (54%) 13.3 17.1%

Adult females 25 (32%) 29.4 37.6%

Subad. females 4 (5%) 2.6 3.4%
Ad. + Subad. females 29 (37%) 32.0 41.0%

Juveniles 7 (9%) 32.7 41.9%

Total 78 (100%)
Ad. = adult, Subad. = subadult. For statistical analysis see text.

*) The contribution of each sex/age class to the troop's composition is expressed as the

mean daily percentage, since troop composition varied over the course of the study. This

value was used to calculate the number of injuries expected in each sex/age class.

(Table 3). The distribution of injuries departed significantly from that ex­
pected according to the respective proportions of adult and subadult males,
adult and subadult females, and juveniles in the troop (X 2 = 82.01, df = 2,
p < 0.0001). Adult and subadult male yellow baboons had proportionally
more visible wounds than females and juveniles (Table 3). The difference
in wounding incidence between males and females was still significant
when juveniles were excluded from the analysis (X 2 = 30.26, df = 1,
p < 0.0001). The exclusion of juveniles is justified on the grounds that
observed wound frequencies are underestimated in this age class, since a
probably large proportion of wounds are lethal (e.g. Altmann, 1980, p. 203;
personal observation), and thus lead to the disappearance of the juvenile
before the wound is noticed.
Nineteen (83%) of 23 adult males belonging to the study troops sustained
at least one injury attributed to agonistic encounters during the 16 months of
the study. Since the residence time of the study males varied considerably
(range 1 day to 5.5 years), the rate as victim in damaging fights was
calculated from the observation time on each individual and its number of
452 CARLOS DREWS

TABLE 4. Individual rates of damaging fights resulting in injury to six adult

and two subadult males in troop Viramba 2

Male vtctim!' Rank D ays2) Injuries Fights 3 ) Days Fights Injuries

per fight per day per fight

HUZ alpha 213 9 3 71 0.014 3.0

ADU beta 213 6 6 35 0.028 1.0
USI lr 163 8 8 20 0.049 1.0
NDU lr 40 0 0
NOV lr 65 0 0
WIM Jr 61 1 1 61 0.016 1.0
sUTE lr 194 4 3 64 0.016 1.3
sNAF lr 213 2 2 106 0.009 1.0

Adult males: Mean days/fight = 46.8, SD = 23.4 (N = 4)

alpha = top ranking throughont study period, beta = second ranking throughout study
period, lr = low ranking (rank anywhere below beta).
1) Only males observed for over one month are listed. These males were not necessarily
simultaneous residents.
2) Days = days covered by observation regime, assuming that on any given observation day
all injuries of the preceding six days would have been detected, given that the minimum
healing time recorded was eight days (Table 7).
3) All fights in the table are damaging. Separate injuries were attributed to the same fight
when dated equally, thus data in the table show independent wounding events.

independent wounding events (Table 4). That calculation took into account
the fact that wounds were still recognizable with considerable certainty up
to six days after the damaging fight (see legend for Table 4). Crudely,
on average an adult male is injured in a fight with another male every
1.5 months, but variation between males is considerable. The alpha male
experienced the highest number of wounds per damaging fight but its rate
of damaging fights was comparatively low. One low ranking male showed
the highest rate of damaging fights of the sample, acting as the scapegoat
among higher ranking males who initiated most contests.
The proportion of aggressive interactions between males which results
in injury can be calculated only by indirect means, given the small sample
size of wound inflictions witnessed during focal animal samples. The
median proportion of aggressive interactions among 20 male dyads in the
main study troop was 10.3%. These were interactions including at least
one aggressive element (i.e. threats, attacks, chases and/or fights). Out of
 INJURIES IN MALE BABOONS 453

91 aggressive interactions between males, only 5.5% resulted in escalation

with high potential for injury (i.e. face-to-face combat, bites, grappling
and/or contact during chases). This corresponds to 1% of all male-male
interactions (N = 662). A male yellow baboon interacts with other males
on average 2.4 timeslh (Drews, 1993b, p. 134). Thus, a male participates in
escalation to potentially damaging levels about 0.024 timeslh, or once every
3.5 days (42 daylight hours). If a male is on average injured once every
47 days (Table 4), then the incidence of injury is about 13 times smaller
than the frequency of escalated fights. This crude calculation suggests that
about 8% of escalated contests result in injury, i.e. 0.4% of all aggressive
interactions or 0.04% of all interactions between males.

Bodily location of injuries

In male yellow baboons, injuries concentrated on the anterior body parts,

particularly on the head (Table 5), as expected from their face-to-face fight­
ing pattern. Wounds on the anterior body parts attributed to agonistic en­
counters were significantly biased towards the right side of the body (66%)
of adult and subadult males (Fig. 1). The wounds of twenty adult males and
two subadults were pooled in this analysis, with no single male contributing
with more than 12% to the sample.
In contrast to the anterior prevalence of wounds in males, most female
injuries were found on their posterior body parts (Table 6, X2 = 34.17,
df = 1, p < 0.0001). The respective proportions of wounds on swellings

TABLE 5. Bodily distribution of injuries resulting from aggression between

adult male baboons in Mikumi

Body part Number of injuries

Head 42
Neck, shoulders and chest 25
Arms 17
Trunk o
Pelvic region 4
Legs II
Tail o
Total 99
454 CARLOS DREWS

RIGHT LEFT

44 23
2 - tailed Binomial p<O.O 15

Fig. 1. Laterality of wounds in adult and subadult male baboons, illustrated in a frontal,
schematic view of a male baboon showing the head, neck, shoulders, upper arms and chest.
Incidence of wounds on lower arms are excluded because these could be moved to the
opposite side during fights and could, therefore, obscure a laterality effect. For details of
sample see text.

TABLE 6. Anterior and posterior bodily distribution of injuries attributed to

agonistic encounters in adult males, adult and subadult females in Mikumi

Body region"? Females Males

Anterior:
Head, Neck, Shoulders, Arms 11 (32%) 84 (85%)
Posterior:
Pelvis, Tail, Legs 5 (15%) 15 (15%)
Sexual swelling 18 (53%)

Total 34 (100%) 99 (100%)

For statistical analysis see text.

*) No injuries were recorded on the trunk (dorsally or ventrally) of either males or females.

 INJURIES IN MALE BABOONS 455

attributed to agonistic interactions (personal observation) and to scratches

against vegetation and alike is unknown. At the sample size used above,
the sex difference in bodily distribution of injuries due to aggression is
statistically significant if a minimum of 11 % of swelling injuries resulted
from aggression (X 2 = 5.62, df = 1, p < 0.02).
Males were sometimes injured on their hindquarters (Table 5). Highest
ranking males are not immune to injuries on their hindquarters. Three
alpha males in different troops received four of the 14 posterior injuries
recorded. Posterior injury of alpha males could happen for instance when
running away from a coalition attack or when pursued by a screaming
subordinate (see Packer, 1979b).

Healing time

A remarkable healing capacity and ability to survive severe wounds is often

commented on for monkeys (e.g. Schultz, 1969, p. 114) and baboons in
particular (Bramblett, 1967). The severity of an injury and the duration of
impairments associated with it are reflected in the healing time. Typically,
fresh wounds dried up within 24 h of the wounding incident. All injuries
in which the healing process was followed up were less than one week old
when first noticed. Data for healing time are summarized in Table 7 and
Fig. 2. The median healing time of injuries from the day that they were
first noticed was about three weeks. There is a slight underestimate in
healing time due to the fact that, on average, wounds were a few days old
when first noticed. This slight bias, however, is compensated by the fact
that due to the irregularity of visits to the troop, the wound was usually
recorded as healed a few days after it had in fact healed.

TABLE 7. Median and range of healing times in days for baboon injuries

N Median Minimum Maximum

Total sample» 35 23.0 8 120

Small injuries « 5 em) IS 14.0 8 114
Large injuries (> 5 em) II 24.0 10 120

*) The sample is composed of 24 cuts, 4 punctures, I tear, I case of limping, I bruise, and
3 injuries of unspecified shape.
456 CARLOS DREWS

12

10

>­ 8
N=35
o
Z
W
::::> 6
a
w
0:::
LL 4

o
10 20 30 40 50 50 70 80 90 100110120130

DAYS TO HEAL

Fig. 2. Frequency histogram of healing times of 35 baboon injuries. For details of sample
see Table 7.

The length of a wound is an insufficient indicator of its severity. The

difference in healing time between small and large wounds (Table 7) was
not statistically significant (Mann-Whitney test, Z = -1.461, N, = 15,
N 2 = 11, p = 0.14). This difference was not significant either when only
cut wounds were considered. The two wounds which took longest to heal
were a 3 em vertical slash splitting the upper lip (114 days) and a 15 em
vertical slash deep in the chest (120 days). Depth of the wound is probably
an important determinant of healing time, but it was not recorded in this
study.

Damaging fights and male immigrations

Escalated fights could be associated with the recent arrival of newcomer

males to a troop, when knowledge about the competitive abilities of op­
ponents is lacking. The null hypothesis that the interval from male im­
migration to a fight resulting in injury among troop males is the product
 INJURIES IN MALE BABOONS 457

20

18

16

14
>­
U 12 l-
z
w
::::l 10 I-
a
w 8
l-
0:::
u,

6 I-

4 l-

2 I-

o
o 15 30 45
I I
60 75 90
Il nil
105 120 135 150

DAYS SINCE IMMIGRATION

Fig. 3. Observed (solid bars) and expected (open bars) frequency distribution of intervals
from immigration to the first fight resulting in injury. Expected values correspond to a
negative exponential distribution and did not depart significantly from the observations
(see text).

of chance, would predict a negative exponential distribution of interval

frequencies (Fig. 3). Expected frequencies n were calculated from

n = NWe-t/T
T
for each observed interval length t, where N is the total sample size (47 in­
tervals), w is the width of the histogram categories (15 days) and T is
the mean interval length observed (T = 46.2 days). The distributions
of observed and expected frequencies of interval lengths did not differ
significantly (Kolmogorov-Smimov Goodness-of-fit test, D max = 0.1477,
N = 47, p > 0.2, Fig. 3), thus failing to reject the null hypothesis that time
since immigration and the occurrence of damaging fights are not causally
related.
458 CARLOS DREWS

Number of wounds per damaging fight

In this section, wounds per fight refers to the number of wounds suffered
by one animal in a single fight, while fight refers to combats resulting in
injury. Hence, the minimum possible value is one, since this analysis is
based on wound records. Wounds on an animal, that were dated back to
the same day are assumed to have happened during the same fight. In 95%
of the fights resulting in injury, the number of wounds sustained by one
individual did not exceed four (N = 67). The mode and median number of
wounds per fight was one (74% of all cases) and the maximum six (Fig. 4).
The null hypothesis that the number of wounds per fight is the product
of chance would predict a negative exponential distribution of value fre­
quencies (Fig. 4). Expected frequencies n for each observed count x were

60 ­

50

40
>­
0
z
W
:J 30
0
W
0::
1.L.
20

10 ­

0
0 2 3 4 5 6 7

WOUNDS PER FIGHT

Fig. 4. Observed (solid bars) and expected (open bars) frequency distribution of number
of wounds, which resulted from fights causing injury. Expected values correspond to a
negative exponential distribution and did not depart significantly from the observations
(see text).
 INJURIES IN MALE BABOONS 459

calculated from
_ N -x/Ai
n- Me ,

where N is the total sample size (69 fights) and M is the mean number
of wounds per fight observed (M = 1.46). The distributions of observed
and expected frequencies of wounds per fight did not differ significantly
(Kolmogorov-Smirnov Goodness-of-fit test D max = 0.0956, N = 67,
p> 0.2).
The average number of wounds did not differ significantly between dam­
aging fights on days in which at least one proceptive female was consorted
in the troop and those on days without consorted females (Mann-Whitney
test, Z = -1.059, N I = 25, N2 = 22, P = 0.29). The proportion of
damaging fights resulting in more than one wound (i.e. above the pop­
ulation median) did not differ significantly either between these groups
(X 2 = 1.175, df = 1, p = 0.28).
The number of wounds per damaging fight was not related in a simple
way to the immigration of males into the troop. Damaging fights soon
after the immigration did not yield significantly more wounds than later
damaging combats (Spearman's r = -0.117, N = 47, p > 0.2). Although
days during which proceptive females were consorted are unlikely to be a
confounding element in this analysis (see above), such days were removed
from the sample in an additional test but the correlation between wounds per
fight and days since last immigration was still not significant (Spearman's
r = -0.136, N = 19, p > 0.5).
Although the time since immigration was not related to the number of
wounds per damaging fight it should be noted that four independent cases
of unwitnessed fights resulting in more than three wounds to one contestant
were associated with recent male immigrations. These cases represent the
upper end (6% of the sample) of the frequency distribution of wounds
per fight, thus suggesting that the arrival of newcomer males leads to the
highest escalation levels during damaging fights.

Implications of injury

Feeding impairment
A male whose upper lip was slashed vertically, was unable to suck in water,
the baboon's usual way of drinking, and had to resort to the more time
460 CARLOS DREWS

consuming lapping. The cheek pouch of another male was sliced open in a
fight. Subsequently, food ingested (fruits of Strychnos madagascariensis)
was seen coming out through the wound.

Drop in dominance rank

A one-step drop in dominance rank associated with injury was observed
in the main study troop in Mikumi. The change in dominance status took
place between the subadult male sUTE and the next ranking dominant male
USI after the latter was noticed limping badly. Comparisons were made
between interactions during the 30 days prior to the injury and the 30 days
thereafter. Whereas USI won 43 out of 44 agonistic interactions against
sUTE during the 30 days prior to the injury, sUTE won all 18 interactions
recorded during the 30 days following the date on which USI sustained
a leg injury (X 2 = 57.4, df = 1, p < 0.0001). All of 37 interactions
recorded in focal samples prior to the injury were initiated by USI while
eleven interactions recorded thereafter were invariably initiated by sUTE
(X 2 = 48.0, df = 1, p < 0.0001). In this case it is unclear whether
the injury was cause or effect of the change in dominance status because
the wounding event was not observed. sUTE left the troop before USI's
leg had healed, so, it is not known whether USI would have regained his
dominant status over sUTE upon recovery.

Changes in behaviour upon injury

The rate of interactions initiated by adult male USI towards the lower
ranking subadult male sNAF decreased significantly after the date of his
leg injury (Binomial test, N = 14, p < 0.05), despite the fact that USI
maintained his dominant status over that male after sustaining the injury.
As a control, the alpha male HUZ and beta male ADU did not change
significantly their rate of interactions towards sNAF during the same period.
There was a significant tendency for male USI to appear in the periphery
of the troop more often when he was limping than before (X 2 = 4.76,
df = 1, p < 0.05). There was no such a significant tendency among
control males HUZ and ADU. USI's time budget, however, was not affected
significantly by his severe limping. The means for time resting, stationary
feeding, feeding on the move, moving, socializing and time spent on the
ground did not differ significantly between the periods before and after
 INJURIES IN MALE BABOONS 461

the injury. Neither differed the distance of USI to the nearest adult male
significantly between these periods.

Discussion
Patterns of injury

The vast majority of wounds in male baboons are canine slashes resulting
from intraspecific aggression. Fractures and other cases of limping can at
least in part be attributed to agonistic encounters resulting in falls from trees
(also Bramblett, 1967). Drops from as high as 12 m during chases (but none
resulting in injury) were observed in this study. In some cases, while the
screaming victim was hanging from the thinnest twigs the attacker shaked
the branch vigorously with a hand, inducing the fall. Harding (1980) saw
an olive baboon male fall off a cliff upon being pursued by another male.
The victim was stunned for 10 min upon impact. Bramblett (1967) reports
from a sample of 61 baboon skeletons, that most fractures occur in the
phalanges of fingers and toes, but fractures on limbs, vertebrae, clavicles,
ribs and skull (mainly on the face) are also common.
In agreement with this study, in olive baboons the wounds on females
(and aggressive bites between juveniles of both sexes) concentrate on the
back and tail, whereas the wounds on adult males are mostly on anterior
body parts, particularly on the face (Owens, 1975; also rhesus macaques
Macaca mulatta: Hausfater, 1972). The proportion of wounds on the
head of male yellow baboons (42%) was similar to that found in olive ba­
boon males (51 %, Owens, 1975). In contrast to yellow baboons, however,
wounds on the back and body flanks were not uncommon in adult male
olive baboons (Owens, 1975). The location of injuries in male hamadryas
baboons (Papio hamadryas) differs from that of yellow baboons, in that
most injuries are inflicted on the hands and forearms despite the fact that
most bites in that species are aimed at the neck (Kummer, 1968, p. 49).
The high incidence of wounds on the head of yellow baboon males could
suggest that canine blows are targeted mainly at the head. This distribu­
tion, however, results more likely from the fact that blows are fended off
with the muzzle, thus exposing the head to injury. In aggressive play, male
juvenile baboons (but not female juveniles) bite the neck of the opponent
more than any other region of the body (Owens, 1975), suggesting that
462 CARLOS DREWS

adults may as well be targeting their blows principally at the neck region.
Analogously, although fights in most ungulates involve frequent head to
head contact, blows are typically aimed at the neck, flanks and haunches
(Geist, 1966, 1986).
There was a right side bias in the lateral distribution of wounds on
male baboons. Behavioural side biases in animals have received attention
in research from the perspective of brain lateralization theory (e.g. Ward
& Hopkins, 1993). Most reports and experiments have addressed asym­
metries in hand use in primates (review in Marchant & McGrew, 1991;
Bradshaw & Rogers, 1993; Ward & Hopkins, 1993; yellow baboons: Post
et al., 1980; Guinea baboons Papio papio: Fagot, 1993; Vauclair & Fagot,
1993) or derivatives thereof, such as the left nipple preference in suckling
chimpanzees Pan troglodytes (Nishida, 1993). Overall, very few cases
are known of limb independent behavioural asymmetries, such as that of
successful slashes during baboon combat reported here. This lateralization
of injuries could be related to an asymmetrical use of the field of vision
during contests, in the way hypothesized for gelada baboons Theropithecus
gelada (Casperd & Dunbar, in prep.).
In contrast to antlers or horns (Geist, 1966), canines are an inefficient
defensive organ when used to parry the opponent's slashes. In baboons
simultaneous injury is likely because at the instant of striking, the attacker
exposes itself to counter-attack, but also because the higher the intensity
of attack the greater the likelihood of retaliation and retaliation increases
the likelihood of intensification of further attacks (game-theory: Maynard­
Smith & Price, 1973; polecats Mustela sp.: Poole, 1973; rhesus macaques:
Bernstein & Gordon, 1974). In fights between olive baboon males, mutual
biting is more common than non-mutual biting (Owens, 1975). Thus, both
the winner and the loser are likely to be injured (also Smuts, 1985, p. 86 and
p. 152), regardless of differences in competitive ability. This is an important
reason for both contestants to avoid escalated fights. Accordingly, in male
yellow baboons only 5.5% of aggressive contests led to high risk escalation
and about 8% thereof resulted in wounds, whereas in red deer stags, a
species with effective defensive organs, 24% of aggressive contests led
to fights (Clutton-Brock & Albon, 1979) and only 2% of fights resulted
in injury (Clutton-Brock et al., 1979). Similarly, severe wounding was
observed in only 1.3% of fights between male white-eared kob Kobus kob
 INJURIES IN MALE BABOONS 463

leucotis (Fryxell, 1987). Consistent with the nature of the weaponry, in

the goat antelope Oreamnos, a species with short, straight and sharp horns,
overt fighting is relatively rare, probably because the risk of injury to
both participants is high (Geist, 1966). In male toque monkeys (Macaca
sinica), another primate with large canines, only 0.6% of threats resulted in
actual body contact (Dittus, 1977). Fights between elephant seal Mirounga
angustirostris bulls, which almost invariably lead to injury, represent only
about 2% of their aggressive conflicts (Le Boeuf, 1971).
Baboon contests involve ritualized aggressive signals, such as eye-lid
displays, ground slapping, audible chewing or teeth grinding, and canine
displays, which probably contribute to an assessment of the opponent with­
out resorting to damaging escalation levels (Smuts, 1987; pers. obs.). The
incidence of injuries shows that contests between male baboons are not
fully ritualized, however. Among species capable of inflicting injury, the
evolution of wholly symbolic conflict is not possible because a mutant that
employed physical attack would have a reproductive advantage (Maynard­
Smith & Price, 1973). Similar to other species mentioned above, however,
the overall frequency of damaging fights between male yellow baboons
is low if expressed as a proportion of all aggressive incidents (less than
1%; also olive baboons: Owens, 1975). Conversely, damaging fights be­
tween male yellow baboons are relatively common, if frequency is based
on time units, for each male is likely to be injured on several occasions
each year. This wounding rate may not be uniquely high among primates.
Each olive baboon female in Gilgil (Kenya) can be expected to receive
at least one serious wound every year, almost invariably inflicted by adult
males (Smuts, 1985, p. 87 ff). The individual rate of fights calculated for
rhesus monkeys was one fight every 26 to 30 days (Symons, 1978, p. 166,
using data from Southwick et al., 1974). In a squirrel monkey (Saimiri
sp.) study all 15 males of the study troop were wounded at some point
(C. Mitchell personal communication to Baldwin, 1992). For references to
wounds in other primate species see Smuts (1987, Table 32.2 therein). Al­
though wounding rates of some ungulates can be similarly high (see Geist,
1986), red deer stags, for example, are wounded in fights only once in a
lifetime (Clutton-Brock et al., 1979).
The incidence of wounds was higher in male than in female yellow
baboons, a sex difference found also in olive baboons (Smuts, 1987),
464 CARLOS DREWS

macaques and other primates (Wilson & Boelkins, 1970; Hausfater, 1972;
Drickamer, 1975; Smuts, 1987). There is no sex difference in the overall
frequency of fractures in baboons (Bramblett, 1967). Virtually all baboons
older than 13 years show fractures (Bramblett, 1967). It should be noticed,
however, that fractures are only a small proportion of the sample which
revealed a difference between the sexes in this study. Overall, therefore,
baboons fit the pattern of several animal taxa where severe aggression,
based on wound frequencies, is more common in males than females (e.g.
crustaceans: Jones, 1980; rodents: Christian, 1971; Rose & Gaines, 1976;
Rose, 1979; ungulates: Geist, 1986; primates: Smuts, 1987).
The median healing time for male baboon injuries was about three weeks.
Smuts (1985, p. 93) reported an 8 ern gash inflicted by a male on the arm
of a female which took two months to heal, the arm not being used for
over one week. Limping from gashes can take several months (Packer,
1979b) and up to a year to heal when recovery is impeded by infection
(Altmann, 1980, p. 210). Intensive grooming of wounds by other troop
members was not uncommon during this study (pers. obs.). The wounds
of adult males were usually groomed by females. This behaviour along
with self-grooming and licking may foster the healing process (Bramblett,
1967). Clean cuts are reported to heal rapidly while dirty wounds in which
a large pocket of loose skin collects debris are often fatal, partly because
adequate cleaning through grooming is not possible (Bramblett, 1967).

Contexts offights resulting in injury

Hausfater (1975, p. 135 if) found that presence of proceptive females in­
creases the incidence of wounds in male baboons in Amboseli National
Park. Here I analysed whether damaging fights in the presence of pro­
ceptive females resulted in more wounds than damaging fights in other
contexts. This was not the case. The likelihood of injury and the number
of wounds sustained in each damaging fight between male baboons were
not related in a simple way to the time since male immigrations either.
The null hypothesis that the number of wounds per fight merely reflects
the result of chance events during the contest, instead of reflecting esca­
lation level, could not be rejected. It is noteworthy, nevertheless, that the
four damaging fights yielding the highest number of injuries concerned re­
cent immigrations or attempts to immigrate by young males in their prime.
 INJURIES IN MALE BABOONS 465

This suggests that the number of wounds per damaging fight could reflect
escalation level at least in some contexts.
Young adult males typically attempt to gain alpha status upon immigra­
tion (Altmann et al., 1988; Noe & Sluijter, 1990). Alpha rank is the most
valuable social resource of young adult males, for their reproductive suc­
cess during this stage of their life depends entirely on holding top rank as
long as possible (Altmann et al., 1988; Noe & Sluijter, 1990). Challenges
for alpha rank are therefore very likely to result in high levels of escala­
tion, irrespectively of the immediate presence of proceptive females, since
the benefits at stake are very high to both contenders. In addition, fights
between young adult males are more likely to result in injury than contests
with older foes partly because the canines of the latter are usually bro­
ken, worn, or both (Packer, 1979b; Smuts, 1985, p. 146). Also, while the
frequency and severity of fighting over rank is expected to be minimized
among close relatives (West-Eberhard, 1975), adult male baboons in a troop
are unlikely to be closely related (Packer, 1979a) and thus, their aggression
is usually not constrained by kinship. Hostility towards immigrant males
sometimes even results in the eviction of the newcomer (Drews, 1993b,
p. 312 jj). A similar pattern is found in other multi-male primate species,
in which violent fights and severe wounds in adult males (and females)
are associated with male immigrations (e.g. rhesus macaques: Lindburg,
1969; Vessey, 1971; squirrel monkeys: Baldwin, 1968). In captive rhesus
macaques a newly arrived male may effectively join a group with high
initial status if he can inflict a few quick injuries to the resident males
(Bernstein et al., 1974).
Injury in male yellow baboons was associated with immediate compe­
tition over proceptive females in three of the ten witnessed incidents of
wounding. Injury to at least one of the contestants was caused in three
(23 %) out of 13 aggressive consort take-overs witnessed, a proportion
similar to that of male gelada baboons fighting over groups of females,
whereby four (25%) out of 16 fights led to severe injury (Dunbar, 1984).
Among the olive baboons studied by Bercovitch (1988) this proportion
was smaller: four (9%) out of 46 aggressive take-overs included wound­
ing. None of the fights observed by Kummer (1968, p. 50) between male
hamadryas baboons was over proceptive females, but rather over long term
possession of a female in the harem. Competition over mating opportuni­
466 CARLOS DREWS

ties is probably a relatively common context of damaging fights in other

primates as well. Male macaques are wounded more frequently during
the mating than during the non-mating season (Vandenbergh & Vessey,
1968; Wilson & Boelkins, 1970; Dittus, 1977; review in Symons, 1978,
p. 173, 174), and 87% of male deaths (but not of females) occur during
the mating season in Cayo Santiago (Wilson & Boelkins, 1970). Simi­
larly, in microtine rodents (Turner & Iverson, 1973; Rose & Gaines, 1976;
Rose, 1979), pinnipeds (e.g. Le Boeuf, 1971) and ungulates (e.g. Wilkinson
& Shank, 1977; Clutton-Brock et al., 1979,1982, p. 132jf), high levels
of wounding in males are associated with the reproductive period. The
proportion of damaging fights among combats associated with immediate
reproductive competition is higher in baboons than in ungulates for which
such proportions have been reported (less than 3%: Clutton-Brock et al.,
1979; Fryxell, 1987).
Challenges to the resident alpha male and fights over proceptive females
are the contexts which most directly affect the reproductive success of
male baboons and, therefore, escalation risking injury may payoff for
both contestants in such situations (Enquist & Leimar, 1990). Fights over
unusual foods (also Ransom, 1981), intertroop encounters, female defense,
redirected aggression and some fights unrelated to any obvious resource
also resulted in injury, however. The relative contributions of these contexts
to the incidence of male wounds cannot be estimated with confidence from
the small sample of ten witnessed inflictions of injury. This diversity of
contexts can obscure a simple unifactorial effect in analyses of determinants
of wounding incidence.

Implications of injury to male baboons

Wounds sustained in fights between male baboons can imply loss of eye­
sight (Packer, 1979b, two cases), bacterial infection (Bramblett, 1967; Alt­
mann, 1980, p. 210 and p. 214) and increased vulnerability to parasites.
Feeding impairments result from injury in the mouth region (P.c. Lee,
pers. cornm.; this study). Wounds on forelimbs can also impair feeding
by constraining the rate at which food items are processed, since baboons
use their hands to pick these items. Also, injury resulting in limping can
hinder low ranking male baboons from their typical running ahead of the
troop to utilize for some minutes a good feeding site, prior to the arrival
 INJURIES IN MALE BABOONS 467

of higher ranking males (pers. obs.). Such locomotory handicaps imply

additional energy expenditure, impaired flight response during predator at­
tack and impaired climbing ability limiting efficient access to food and
safe retreats. Furthermore, injury can affect the social activity of male
baboons, as shown in this study in the form of a reduction in the rate of
interactions initiated. Similarly, injured olive baboon males generally avoid
interacting with other males (Popp, 1978, p. 100; Harding, 1980; Ransom,
1981), whereas other troop males seek out repeatedly and aggressively the
handicapped peer (Harding, 1980). As a consequence, some injured males
become peripheral or emigrate temporarily (Harding, 1980; Collins, 1981,
p. 336; this study).
The costs of non-lethal injury in male baboons can be substantial. Such
injury, particularly when impairing locomotion, can affect fitness directly
by reducing the victim's competitive ability, causing drops in dominance
rank and jeopardizing its mating success. Drops in dominance associated
with injury to males as observed during this study have been reported from
another population of yellow baboons (Altmann, 1980, p. 210) and for olive
baboons (Popp, 1978, p. 100; Ransom, ] 981). A drop in dominance rank
implies for the victim an increment in the number of potential supplanters
and aggressors. Injuries have been found to reduce the reproductive activity
of male baboons in at least six independent cases (Packer, 1979b; Collins,
]981, p. 336; Smuts, 1985, p. 133). In other male primates, injury or
illness also results sometimes in drops in dominance (e.g. rhesus macaques:
Kaufmann, 1967; Vandenbergh, 1967; chimpanzees: Bygott, 1979) and in
a reduction in mating activity (e.g. squirrel monkeys: C. Mitchell pers.
comm. to Baldwin, 1992; rhesus macaques: Conaway & Koford, 1965;
Loy, 1971).
Fights between male baboons can result in death to a combatant. Ransom
(1981) reported that an old, adult olive baboon male in Gombe National
Park sustained a slash wound on his inner thigh in a fight with another
male over bananas. The femoral artery was severed and the male bled
slowly to death. Another male at the same site is suspected to have died
in a fight over a proceptive female (1. Goodall pers. comm. to Packer,
1979b). In Amboseli (Kenya), a high ranking male suffered a severe leg
injury probably inflicted by an uprising natal male (Altmann, 1980, p. 205
and p. 214). The puncture wounds became badly infected until a month
468 CARLOS DREWS

later the male could hardly move and disappeared overnight. Two olive
baboon males were killed as single combatants against coalition attacks
on independent occasions in the Masai Mara, Kenya (Popp, 1978, p. 96).
Popp inferred that approximately 20-30% of all adult males in this area
die directly from injuries received from other males and that almost all of
these lethal injuries are inflicted by coalitions. Bercovitch (1988), however,
conjectures that baboon male coalitions in Gilgil, Kenya, are less likely to
result in injury than dyadic fights, both on an absolute and a per capita
risk basis. Canine gashes sustained during fights were the major cause of
mortality in male chacma baboons Papio ursinus belonging to a desert troop
(Brian, 1992). In that study, four males died and two disappeared after
sustaining serious wounds during fights. These lethal fights occurred during
male immigration and competition over reproductive females (Brian, 1992).
Lethal injuries sustained in fights between males are known from other
primates as well (e.g. rhesus macaques: Lindburg, 1971; chimpanzees:
Goodall, 1986, p. 503 jf).
Across anthropoid primates there is a positive association between fre­
quency and intensity of escalated aggression between males and the degree
of canine dimorphism in the species (Plavcan & van Schaik, 1992). It
is argued that selection has favoured large canines in male primates as a
means to enhance the chances of winning fights (also Martin et al., 1994).
This argument rests on the assumption that effective use of canines during
escalated contests leads to injuries resulting in costs to the opponent. The
evidence reviewed here shows that the handicaps resulting from damaging
fights in male baboons and other primates can be substantial.
In general, the balance between value of resource contested and future
reproductive prospects is a major determinant of the likelihood of severe
injury in fights (Enquist & Leimar, 1990). Although the implications of
injury sustained in fights between male baboons can be considerable, it
should be noticed that most wounds were small, healed well and did not
seem to affect fitness in any direct way, for not uncommonly the victim of
wounding was the winner of the fight (also in elephant seals: Le Boeuf,
1971). Thus, while the risk of injury attached to escalated contests be­
tween males can be a more or less constant value, the costs resulting from
injury can vary considerably. It is likely that chance determines to some
extent the severity of canine slashes inflicted in a fight, i.e. the depth of
 INJURIES IN MALE BABOONS 469

the wound, organs affected, etc. Consequently, the decision to escalate a

fight should incorporate a low predictability of injury costs in damaging
encounters.
In conclusion, the costs potentially associated with escalated aggression
seem high enough to exert strong selective pressure to both possess highly
damaging weaponry and to avoid escalated confrontations. Avoidance of
escalated confrontations is seen in the very low percentage of such es­
calation from the total of aggressive contests between males. Although
escalation is generally avoided, the high interaction rate between male ba­
boons results in a relatively high risk of injury. The intra-sexual selective
pressure acting on males is reflected in the fact that male baboons are
repeatedly injured in fights during their lifetime. The implications of in­
jury listed above are the basis to understand patterns of contest avoidance,
peaceful conflict resolution, punishment (Clutton-Brock & Parker, 1995),
as well as the dynamics of dominance relationships between male baboons
and possibly other primates with similar patterns of reproductive competi­
tion.

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