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Contexts and Patterns of Injuries in Male Baboons - Drews 1996

Contexts and Patterns of Injuries in Male Baboons - Drews 1996

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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 damagingfights in order to provide a framework for analyses of conflict resolution and dynamicsof agonistic competition in primates. The vast majority of wounds were canine slashesresulting from intraspecific face-to-face combat. Wounds were more common in malesthan 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 conflictsrepresented 10% of all interactions between males. Less than 1% of aggressive contestsled to injury. The individual rate of injury from fights with other males was on averageonce 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 thepresence 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 andTechnology, 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 hostedthe author at the respective study sites. The field assistance of A. Sindimwo in Gombe andof C. Kidung'ho, W. Marwa and A. Njalale in Mikumi was invaluable. I am most gratefulto A. Schmidl-Drews, who collected half of the data for this study. Previous versions ofthis 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 fundedby 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. Thepreparation 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 immigrateby adult males in their prime. Contexts of male injury observed during infliction includechallenges 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 fightunrelated to any obvious resource. This study and anecdotal reports from the literature pointat various implications of injury to male baboons, including physical impairments whichcan constrain feeding efficiency, limit access to resting sites and safe retreats, cause a dropin dominance rank, jeopardize mating success and even result in death. Severely injuredmales typically reduce interaction rates, retreat to the periphery of the troop or emigratetemporarily. Although most wounds are small and heal well, the potentially high costsof injury probably exercise strong selection pressure on contestants for means of peacefulconflict resolution, given that during fights both baboons risk injury irrespective of theircompetitive abilities.
The
potential fitness consequences of inflicted injury can explain theevolution 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 (seeHuntingford
&
Turner, 1987). Surprisingly little is known about the consequences of wounds resulting from intraspecific fight, given that the riskof injury and its associated costs are a central component in game-theoryanalyses 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 isknown about the non-lethal implications of injury sustained in fight, however.
In
red deer
Cervus elaphus
stags, recovery from injury is associatedwith 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 fromfield studies concerning the pattern, context and implications of lesions.During escalation animals risk to suffer a potential fitness reduction asa consequence of injury. There can be direct costs of injury, such as
 
445
NJURIES IN MALE BABOONS
e.g.
a greater energetic expenditure from limping or the mating opportunities missed
due
to a reduction in competitive ability. In addition there arestochastic costs which
mayor
may
not be incurred, and arise
e.g.
from anincreased risk
of
falling prey to predators in the case
of
a locomotory handicap. In principle, however, damaging fights, defined as contests leading toinjury to any or both participants, need not necessarilly reduce the fitnessof the victim because injury
can
be
minor
and may heal rapidly. Thus,the type
of
injury, its severity
and
its healing time largely determine thekind and magnitude
of
the handicaps attached to it.
The
aims
of
this reportare
(1)
to present field
data
on lesions in male savannah baboons
(Papiocynocephalus),
(2) to bring together the anecdotal information about maleinjuries 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 competitionin primates
(e.g.
Popp
& DeVore, 1979), but studies of fights are missing inthis order, which thus lacks the empirical data required in some models ofaggressive competition. An insight into pattern and context of damagingfights
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 likelyto observe the result of aggression,
i.e.
injuries, than the aggression itself,because fights resulting in injury are brief and relatively rare, whereasinjuries
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, forexample, 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

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