PRIMATES, 43(3): 223-235, July 2002 223

Conflict and Post-conflict Behavior in a Small

Group of Chimpanzees
AGUST1N FUENTES, Notre Dame UniversiO'
NICHOLAS MALONE, University of Oregon
CRICKEqq'E SANZ, Washington University
MEGAN MATHESON, Central Washington UniversiO:
and LORIEN VAUGHAN, San Diego Wild Anhnal Park
ABSTRACT. Chimpanzee research plays a central role in the discussions of conflict negotiation.
Reconciliation, or the attraction and affiliation of former opponents following conflict, has been proposed
as a central element of conflict negotiation in chimpanzees and various other taxa. In an attempt to expand
the database of chimpanzee conflict resolution, conflict and post-conflict behavior were recorded for a
small group of socially housed chimpanzees at the Chimpanzee and Human Communication Institute, at
Central Washington University, Data were collected over six 6-week periods between 1997 and 2000, for a
total of 840 hours of observation, resulting in a substantial post-conflict (PC) and matched control (MC)
data set. The data demonstrate this group' s tendencies to maintain visual contact and closer proximity after
conflicts. Dyadic corrected conciliatory tendencies ranged between 0 - 37.5% and averaged 17.25% across
all dyads. Individual corrected conciliatory tendencies ranged between 5.8 and 32%. The results of this
study combined with recent publications on captive and free-ranging chimpanzee post-conflict behavior
suggest that variation in post-conflict behavior may be important to our understanding of chimpanzee con-
flict negotiation, and may also have implications for the design and management of captive chimpanzee
enclosures and social groups, respectively.
Key Words: Conflict; Post-conflict behavior; Chimpanzee; Pan troglodytes; Reconciliation.
I NTRODUCTI ON
Pri mat ol ogi st s, Ant hropol ogi st s, and Psychol ogi st s have l ong been interested in aggressi ve
behavi or in nonhuman primates. A large body of research has reveal ed a wi de cont i nuum of
confl i ct across all age/ sex cl asses and the integral rol e it holds in the daily lives of hi ghl y social
group-l i vi ng organi sms (DE WAGE, 1993, 2000; MASON MENDOZA, 1993). Our current vi ew
charact eri zes social confl i ct as a component of subtle and compl ex social relationships, and
i mport ant to the dynami c processes i nvol ved in their f or mat i on and mai nt enance (AURELI t~ DE
WAGE, 2000; DE WAAL, 2000).
The current defi ni t i on of reconci l i at i on operat i onal l y defi nes attracted, dispersed, and neutral
pairs (DE WAAL& YOSHIHARA, 1983). DE WAAL and YOSH|HARA suggest ed t hat f ol l owi ng con-
flict, attraction rat her t han dispersal is likely to occur and f or mer opponent s are likely t o interact
affiliatively t hrough cl ose pr oxi mi t y and body cont act . VEENEMA et al. (1994) refi ned the PC
( post - conf l i ct ) - MC ( mat ched cont r ol ) dat a col l ect i on met hodol ogy with the addi t i on o f a correc-
tion fact or that r emoved basel i ne behavi or from post -confl i ct interaction data (see al so
VEENEMA, 2000). This resul t ed in a mor e robust measure called the Cor r ect ed Conci l i at ory
Tendency ( CCT) (at t ract ed-di spersed pai r s/ t ot al number of confl i ct pairs).
To date, a number of post -confl i ct studies have been undert aken across pri mat e t axa
(revi ewed by KAPPEEER & VAN SCHAIK, 1992; DE WAAL, 1989, 1993; AUREL! & DE WAGE,
2000). I n general the results have demonst r at ed a range in post -confl i ct responses, part i cul arl y
224 A. FUENTES et al.
in rates and styles of reconciliation (see KAPPELER & VAN SCHAIK, 1992; DE WAAL, 1993;
AURELI & DE WAAL, 2000 for overviews of the data sets). Other post-conflict behavior patterns
investigated include consolation (affiliative interaction with third party after a conflict) and re-
direction (aggression directed towards other group members) (see AURELI & DE WAAL, 2000).
One of the major findings emerging from the growing number of post-conflict behavior studies
is that of variability in conflict negotiation and resolution (see AURELI & DE WAAL, 2000;
CASTLES et al., 1996; CORDS & KILLEN, 1998; SCHINO et al., 1998).
Although there are a number of hypotheses for why primates tend to reconcile (or not recon-
cile) after conflict have arisen, two have received the most attention. The two primary models
for conflict resolution are the valuable relationship hypothesis (CORDS & AURELI, 1996) and the
uncertainty reduction hypothesis (SILK, 1996). Both of these models find some support from
existing data, although differing interpretations of the data leave many vagaries yet to be settled
(AURELI & SMUCNY, 1998; CORDS ~, AURELI, 1996, 2000; SILK, 1996, 1998, 2000). It is impor-
tant to note that while the former hypothesis speaks to long-term relations within the group and
the latter emphasizes immediate, or "current" behavior, the two are not necessarily exclusive.
Proponents of both models stress the need for continued and expanded data collection across
primates and other taxa to address these issues. Also, there is a clear need for a larger body of
quantitative data with which to assess these and other models for the behavior of organisms
after conflict.
The purpose of this study is to add to the existing body of literature examining post-conflict
behavior, including reconciliation, in captive chimpanzees (ARNOLD & WHITEN, 2001; DE
WAAL, 1987, 1989, 1993; DE WAAL AURELI, 1996; DE WAAL t~ VAN ROOSMALEN, 1979;
PREUSCHOFI" et al., 2002). The five chimpanzees of the Chimpanzee and Human
Communication Institute (CHCI) are a small group, all adults, where we have long-term knowl-
edge of personal histories and behavioral records combined with excellent observational condi-
tions. While four of the five chimpanzees in this group were cross-fostered all have been in
exclusive chimpanzee groups for the last 19 years. This group' s behavioral repertoire has some
elements not common in other captive chimpanzee colonies, however their overall behavioral
profiles are well within general chimpanzee parameters (JENSVOLD et al., 2001; MARTIN et al.,
1999).
Here we report the first four years' data of a long-term project to examine the conflict and
post-conflict behavior in this small group of chimpanzees.
METHODS
SUBJECTS
See Table 1 for biographical information on this group of five adult chimpanzees (Pan
troglodytes). These chimpanzees have been housed together since 1981. They have resided in
their present enclosure at the CHCI since 1993. Four of the chimpanzees were cross-fostered
and acquired American Sign Language (ASL) from their human caregivers (GARDNER et al.,
1989). The fifth (Loulis) acquired his signs from his adopted mother Washoe and the other
chimpanzees (FOuTS et al., 1989).
The chimpanzees' facility consists of an outdoor enclosure, two indoor exercise rooms, and a
night enclosure. The enclosure encompasses 1909 m 2 and offers 4417 m 2 of functional surface
area. The enclosures are furnished with a variety of structural enrichment items. The chim-
panzees are provided three meals a day in the night enclosure. Each day caretakers provided
social and object enrichment to the chimpanzees (FOUTS et al., 1989, 1994).
Conflict and Post-conflict Behavior
Table 1. Participant biographical information.
225
Participant Age in years as of 2001 Sex
Washoe Approxi mat el y 36 Femal e
Moja 30 Femal e
Tatu 27 Femal e
Dar 26 Mal e
Loulis 23 Mal e
PROCEDURE
Data were collected for a total of 840 hours over six 6-week periods [June - August 1997,
June - August 1998, January - February 1999 (1999a), June - August 1999 (1999b), January -
February 2000 (2000a), and June - August 2000 (2000b)]. Dat a were collected from 09:00 to
11:00 and 13:00 to 15:00 each weekday. Dat a were also collected from 12:00 to 13:00 five days
a week in 1997 and three days a week in 1998, 1999, and 2000. Data were collected via focal
follows by two observers during each collection session so that both participants in a conflict
could be accurately recorded.
Dat a collectors recorded: conflict participants' proximity to one another (in 1 m increments),
visual contact (unobscured line of sight between the conflict participants), behavi or exhibited,
partner (if behavi or was interactive), directionality of behavi or (actor, receiver, or mutual), and
presence of ASL at 20-sec intervals for the 10 min post-conflict and matched control periods.
The first affiliative interaction after a conflict was recorded for all post-conflict and matched
control periods. Observers obtained a mi ni mum of 90% inter-observer reliability with an experi-
enced observer recording behavioral contexts, proximity, reciprocity, and the occurrence of
ASL in interaction.
During the 1998, 1999 (a and b), and 2000 (a and b) data collection periods conflict data
were recorded. Dat a collectors recorded the initiator and recipient of a conflict, beginning t i me
of conflict, duration of conflict, level of conflict intensity, and a narrative description of the con-
flict. Dat a collectors achieved 90% inter-observer reliability with an experienced observer
recording the presence of conflict and level of conflict intensity prior to the data collection
period. We defined conflict intensity by four levels. Level 1 consisted of directed threat behav-
ior with no physical contact. Level 2 consisted of aggressive hits or kicks. Level 3 involved
repeated hits/kicks, dragging and/or grappling, and Level 4 consisted of biting and/or physical
contact resulting in observable injury (blood, broken skin, etc.).
A conflict event was defined as an aggressive interaction involving agonistic contact or three
or more agonistic behaviors directed at another chimpanzee. End of a conflict event was defined
as the cessation of aggressive behaviors between conflict participants. However if aggressive
behaviors between conflict participants were observed within 2 min of the end of the original
conflict, this was considered a false start. Observers waited until the cessation of aggressive
behavi or and then recorded post conflict (PC) data. Corresponding matched-control observa-
tions were conducted on the next observation day. These 10-min MC observations began at the
same time of day and involved the same individuals as PC observations (see VEENEMA et al.,
1994).
Reconciliation, consolation, redirection, and opponent proximity were examined by compar-
ing "attracted" and "dispersed" pairs. I f opponents interacted affiliatively earlier, or only, in the
PC then they were t ermed "attracted" and the interaction was scored as reconciliation in the PC.
I f the affiliative interaction was earlier, or only, in the MC period then the opponents were
termed "dispersed" (see VEENEMA, 2000). I f an opponent-third party affiliative interaction was
226 A. FUENTES et al.
recorded in the post-conflict period then the matched control period was examined for oppo-
nent-third party affiliative interaction. If the pair was termed "attracted" (see above) the conso-
lation was considered to have occurred. Inter-opponent proximity was measured as closer
sooner in the PC than the MC (attracted) or closer sooner in the MC than in the PC (dispersed).
Maintenance of visual contact was scored based on the 30 20-sec scans recorded during the PC
and MC periods. Conciliatory tendency was calculated by dividing the attracted pairs by the
total number of conflicts. Corrected conciliatory tendency (CCT) was calculated by subtracting
the dispersed pairs from the attracted pairs and dividing the result by the total number of con-
flicts (VEENEMA, 2000; VEENEMA et al., 1994). Consolation, redirection, and proximity tenden-
cies were calculated in the same manner (with attracted pairs exhibiting the behavior in question
only or first in the PC, dispersed pairs only or first in the MC and neutral pairs not at all).
Consolation and redirection dyads where humans were participants (separated by enclosure bar-
riers) were also included. In calculating the proximity tendency conflict pairs that were deter-
mined to be either attracted or dispersed through social interactions (see CCT calculation
above) were excluded from this independent analysis of proximity. Statistical analyses pre-
sented here were conducted via Chi-square tests, Wilcoxon signed ranks tests and z-tests for
proportions with a significance level of p<. 05.
RESULTS
CONFLICT DATA
A total of 219 conflicts (224 conflict pairs) were examined (1998 - 2000 collection periods).
One hundred and twenty-nine conflicts consisted of aggressive hits or kicks (Level 2). Eighty-
three conflicts consisted of directed threat behavior with no physical contact (Level 1). Twelve
conflicts involved repeated hits/kicks, dragging and/or grappling (Level 3). No conflicts
resulted in serious injury during the data collection periods, however at least three Level 4 con-
flicts occurred during non-data collection periods between 1998 and 2000. The range of conflict
duration was 2 - 120 sec, with an average conflict duration of 15 sec.
In Figure I we depict the distribution of conflict within the chimpanzee' s enclosure. The
majority of conflicts occurred in the East room. The chimpanzees had full access to the outdoor
portion of the enclosure during the summer observation periods, however during the 1999a and
2000a observation periods access was primarily limited to the indoor rooms due to frequent out-
door temperatures below 0C.
POST-CONFLICT AND MATCHED CONTROL DATA
Post-conflict and matched control data were collected for a total of 255 conflicts (262 con-
flict pairs). Table 2 presents the total post-conflict and matched-control data set for the six col-
lection periods between 1997 and 2000. Data are presented for reconciliation, consolation,
redirection, and proximity for both the MC and PC observation periods (see methods for
description of how these measures were calculated). Table 3a presents the number of conflicts
per dyad and the corrected conciliatory tendency for that dyad. This was calculated by subtract-
ing dispersed from attracted conflicts involving a dyad and dividing by the total conflicts for
that dyad. Dyads were significantly more likely to be attracted following conflict than dispersed
(T=3.5, n= 10, p<. 05). The mean corrected conciliatory tendency across all dyads is 17.25 %.
There is a nonsignificant negative correlation (Spearman' s Rho= - 0. 354, n= 10) between con-
Conflict and Post-conflict Behavior
Fi g. 1. Distribution of conflicts at CHCI 1998 - 2000b.
I Night Area 1
West I East
29 ~ 1 6 5
227
f l i ct r at e per dyad and Cor r ect ed Conci l i at or y Tendency. Di f f er ent i nt ensi t y l evel conf l i ct s al so
var i ed by Cor r ect ed Conci l i at or y Tendency.
Tabl e 3b pr esent s i ndi vi dual cor r ect ed conci l i at or y t endenci es, and Tabl e 3c pr esent s t he cor -
r ect ed conci l i at or y t endenci es by age/ s ex cl ass and is c ompa r e d wi t h t he s ame meas ur es pub-
l i shed by PREUSCHOVr et al. (2002). Whi l e t hese c hi mpa nz e e s r econci l ed an aver age o f 17. 5%
o f al l conf l i ct s acr oss dyads , i ndi vi dual dyads r anged in cor r ect ed conci l i at or y t endenci es ( r ange
- 3 - 37. 5%) .
Tabl e 4 pr esent s t he resul t s of t hi s st udy al ong wi t h t he ot her publ i s hed c hi mpa nz e e post -
conf l i ct dat a.
Whi l e f r equency of par t i ci pat i on in conf l i ct var i ed, pr opor t i onal par t i ci pat i on r e ma i ne d f ai r l y
cons i s t ent t hr oughout t he study. Loul i s , t he younge s t mal e was i nvol ved in t he hi ghest number
o f conf l i ct s (87%). The ot her chi mpanzees par t i ci pat ed l ess f r equent l y: Da r ( 46%) , Wa s h o e
( 32%) , Tat u ( 31%) , and Mo j a (9%).
Tabl e 5 pr esent s t he per cent ages of t ot al conf l i ct s in whi ch r edi r ect i on, cons ol at i on, i nt er-
Tabl e 2. Total post-conflict data 1997 - 2000.*
Conflicts Reconciliation Consolation Redirection Proximity
255 PC 39 PC 105 PC 80 PC 176
(262conflict pairs) MC 15 MC 93 MC 21 MC 83
*Bad observations were recorded for proximity data on two conflicts in 1998 and one conflict in 1999a. PC: In post-
conflict only or first (attracted pairs); MC: in matched-control only or first (dispersed pairs).
Tabl e 3a. Number of conflicts per dyad and Corrected Conciliatory Tendency (CCT) between individual
conflict pairs (attracted-dispersed/total conflicts for that pair).
Dar Loulis Moja Tatu Washoe
Dar - 90 (4.4%) 5 (20%) 13 (30.8%) 3 (-33%)*
Loulis - - 9 (33%) 50 (10%) 78 (3.8%)
Moja - - - 8 (37.5%) 3 (33%)
T a t u . . . . 3 (33%)
Washoe . . . . .
*A negative CCT is reported for this dyad due to a higher frequency of dispersal following the observed conflicts.
228
Tabl e 3b. Corrected Conciliatory Tendencies for each individual.
A. FUENTES et al.
Subject CCT
Dar 8.1%
Loulis 6.4%
Moja 32%
Tatu 17.6%
Washoe 5.8%
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
CCT
Across 17.25%
Dyads
Tabl e 3c. Corrected Conciliatory Tendency by age/sex class dyad at CHCI compared with PREUSCHOFT et
al. (2002) values same age/sex classes.
Age/sex class CHCI (this study) Yerkes (PREUSCHOI~ et al., 2002)
Adult male-Adult male 4.4% NA
Adult male-Adult female 16.3% 45.7%
Adult female-Adult female 34.5% 57.9%
Tabl e 4, Description of reconciliation and Corrected Conciliatory Tendenci es i n captive chimpanzee stud-
ies.
Corrected Conciliatory
Species Location/setting Group size Opponent pairs Reconciliation Tendency
Pan troglodytes Arhem l)
Indoor, 1979 20 150 34.7% -
Outdoor, 1979 20 200 29.5% -
Outdoor, 1981 20 95 26.6% -
Detroit 2)
Indoor/Outdoor, 1994 11 43 49.7% 14.4%
CHCI 3)
Indoor~Outdoor 5 262 25. 1% 17.25%
1997 - 2000
Budongo 4)
Free Ranging, Uganda 51 120 19.2%
Yerkes 5) 16 401 44.8% 41.2%
Outdoor
Pan paniscus San Diego Zoo 6)
Indoor/Outdoor, 1987 6 333 43.8%
Outdoor, 1997 4 179 55.9%
1) DE WAAL VAN ROOSMALEN, 1979; GRIEDE, 1981; 2) BAKER ~. SMUTS, 1994; 3) This study; 4) ARNOLD & WHITEN
2001; 5) PREUSCHOI'q" et al., 2002; 6) DE WAAL, 1987.
o p p o n e n t pr oxi mi t y, and ma i n t e n a n c e of vi s ual c ont a c t we r e r ecor ded f or bot h PC and MC per i -
ods acr os s t he ent i r e dat a c ol l e c t i on sessi on. See MALONE et al. ( 2000) for a det ai l ed des cr i pt i on
of r edi r ect i on and c ons ol a t i on i n t hi s gr oup of c hi mpa nz e e s .
Be c a us e of t he he a vy r e pr e s e nt a t i on of one i ndi vi dua l we a n a l y z e d t he da t a for c ons ol a t i on
and r edi r ect i on e xc l udi ng t he i ndi vi dua l ( Loul i s) as an act or ( and as r e c i pi e nt i n t he cas e of con-
s ol at i on) . Thi s pr oduc e d t he s a me pat t er ns as t he pool ed dat a: no s i gni f i c a nt di f f er ence for con-
s ol at i on PC or MC (z=. 62, n. s. ) and s i gni f i c a nt l y mo r e r e di r e c t i on af t er conf l i ct s ( PC) t han
dur i ng ma t c he d cont r ol s ( MC) ( z=3. 72, p<. 01) .
Pr oxi mi t y ( cl os er s oone r i n PC vs cl os er s oone r i n MC) was al so a n a l y z e d acr oss dyads , t he
Conflict and Post-conflict Behavior 229
Table 5. Group totals for redirection, consolation, overall proximity, and maintenance of visual contact
for both PC and MC expressed as a percentage of total conflicts.
PC MC PC vs MC (chi-square test)
Consolation 40.0% 35.9% NS
Redirection 30.5% 8.0% p < .01
Proximity (closer sooner) 67.2% 31.7% p < .05
Maintenance of visual contact 79.1% 64.6% p < .01
Table 6, Individual and group patterns for behaviors exhibited significantly more often in the PC than in
the MC periods (p < .01).
Dar Loulis Moja Tatu Washoe Group
Agonism 3
Affinitive
Social I, 4 2
Bad
Observation 4 5
Coprophagy 2, 3, 5 2, 3, 5
Display l, 2, 3, 5, 6 2, 3, 5, 6 l, 2, 3, 5, 6
Feeding 6 l, 6 3 6
Groom 3 4 4
Play 4 l l, 4
Object
Manipulation 3 4 2 3, 4, 6 3, 4
Other 5 2 2
Reassurance 3 3
Self groom 3, 5, 6 3, 4, 5 1 3 6 3, 5, 6
Threat 5 1,2,4,5,6 3 1,2,5,6 1,2,4,5,6
Travel 2 1, 4 3 1, 2, 4
1: 1997; 2: 1998; 3: 1999a; 4: 1999b; 5: 2000a; 6: 2000b.
results show that conflict participants were more likely to be closer after a conflict than in a
matched control period (T=0, n=7, p<.05).
Table 6 presents the behaviors that were exhibited significantly more often in the PC period
by individuals and at the group level. A total of 16 behaviors were exhibited significantly more
frequently in the PC period by one or more individuals across one or more of the data collection
sessions, and a total of 11 behaviors were exhibited significantly more often at the group level
during the PC period. Only two behaviors were exhibited in five of the six data collection ses-
sions significantly more often in the PC than the MC by the group (p<.0)). These behaviors
were "Display" and "Threat."
DISCUSSION
CONFLICT DATA
Interactions leading up to conflicts at CHCI include rough play behavior, disputes over a
favored object or location, and territorial displays (directed towards humans). Because of the
small size of this group and its long-term stability our initial expectations were that very few
serious conflicts would occur. The overall conflict rate of approximately one conflict every
three observation hours bears this out. The majority of conflicts were of low to moderate inten-
sity (Levels 1 and 2). However, this does not imply that conflict is not important at the CHCI.
230 A. FUENTES et al.
That measurable behavioral patterns emerged after conflict by group members indicate that
even low intensity conflict may be an important social stimulus (see SCHINO et al., 1998; VAN
SCHAIK AURELI, 2000).
ARNOLD and WHITEN (2001) found a non-significant trend in their free-ranging study of
chimpanzees suggesting that conciliatory tendency is negatively correlated with conflict inten-
sity (they report a mean CT of 36.1% for low intensity conflicts and a mean CCT of 16.7% for
high intensity conflicts). Additionally, PREUSCHOFr et ai. (2002) also report more lower level
than higher level intensity conflicts in their study of captive chimpanzees at Yerkes Regional
Primate Research Center.
During this study one individual was a participant in the vast majority of recorded conflicts.
This fact probably impacts the post-conflict behavior strategies utilized by the other group
members. Loulis is the youngest male and appears to hold a high rank in the group (SANZ et al.,
1995). Interestingly, PREtJSCHOVr et al. (2002) also report that a young male (a subadult in their
case) participated disproportionately in conflicts. However, when we remove Loulis from the
data set we still see the same patterns in consolation and redirection behavior which suggests
that Loulis' impact may not be dramatically changing other chimps' behavior strategies. While
this disproportionate representation of one individual in the conflict data stands out in a small
group it is important to note that conflict participation may vary dramatically by individual in
many situations across many primate groups.
The majority of recorded conflicts occur in the East room. A thick cement wall separates this
room from the inner areas of the research facility wherein 15 - 50 people are frequently moving
about involved in research and classroom activities. It is possible that the location, or structural
aspects, of this area directly influence the high number of conflicts seen here.
POST-CONFLICT DATA
The results presented here share both similarities and differences with the other reports of
chimpanzee post-conflict behavior (ARNOLD & WHITEN, 2001; BAKER & SMUTS, 1994; DE
WAAL, 1987, 1989, 1993; DE WAAL & AtJRELI, 1996; DE WAAL & VAN ROOSMALEN, 1979;
PREUSCHOFr et al., 2002). Our results do reinforce recent reports of variation within and across
primate groups (ARNOLD & WHITEN, 2001; AtJRELI & SMtJCN, 2000; CASTLES et al., 1996;
CORDS & KILLEN, 1998; PREOSCnOVr & VAN SCnA1K, 2000; SCnINO et al., 1998; THIERRV, 2000).
While it is apparent that the dyads observed in this study do reconcile nearly a fifth of their con-
flicts, individual variation across dyads, and the variable post-conflict behavior observed com-
bined with the variation in reported chimpanzee CCTs from other studies suggests that these
chimpanzees may utilize an array of behavior, in addition to reconciliation, when negotiating
conflict.
RECONCILIATION
Unlike previous studies of captive chimpanzees we did not observe a specific set of overt
affiliative behavior occurring after conflicts (see DE WAAL & VAN ROOSMALEN, 1979; DE WAAL,
1989, 1993; PREUSCr~ovr et al., 2002). The one reported free-ranging study of chimpanzee post-
conflict behavior (ARNOLD & WHITEN, 2001) also reports a lack of a clear and discernable char-
acteristic set of reconciliatory behaviors. While embracing was observed as an affiliative
behavior involved in reconciliation, we did not observe frequent kissing, holding out of hand, or
submissive vocalizations. This may be due to the relatively small number of reconciled conflicts
[24 (attracted - dispersed) out of 262 opponent pairs]. Alternatively, this may be because differ-
Conflict and Post-conflict Behavior 231
ent groups of chimpanzees may exhibit differing post-conflict affiliative behavior ("cultural"
differences: see MCGREW, 1998; WHITEN et al., 1999; and differences in age, sex, kin relations,
group history: see ARNOLO & WHITEN, 2001 ).
Given that four of the five chimpanzees in this group were cross-fostered, it is possible that
this has an impact on their post-conflict behavior. However, only a few chimpanzee studies
have been conducted using the PC/MC control method and the correction factor proposed by
VEENEMA et al. (1994) which subtracts the baseline affiliation from post-conflict affiliation.
Therefore it is difficult to make specific comparisons and assertions about baseline chimpanzee
reconciliation behaviors. Two other published studies on captive chimpanzees (Detroit Zoo:
BAKER & SMUTS, 1994; Yerkes: PREUSCHOFF et al., 2002) and one on free-ranging chimpanzees
(Budongo: ARNOLD WHITEN, 2001) provided corrected conciliatory tendencies or contained
sufficient published information allowing us to calculate the corrected conciliatory tendency.
The Detroit Zoo study, the study at Budongo, and this study result in fairly low corrected con-
ciliatory tendencies of between (14.4 and 19.2%). PREUSCHOFT et al. (2002) report a higher CCT
(41.2%). However, one must be extremely careful when reporting group level corrected concil-
iatory tendencies, as they may not accurately reflect the underlying individual variation, and it
is not clear that a mean score effectively represents a trend or group behavioral pattern.
Individuals varied in their rates of reconciliation, most notably by conflict participant. One
female, Moja, engaged in the lowest number of conflicts (25) yet reconciled 32% of them.
Loulis, on the other hand, participated in a high number of conflicts (227) with reconciliation
occurring after very few of them (6.4%). This trend of lower CCT with increasing conflict fre-
quency was also found by PREUSCHOFT et al. (2002). Individuals in the CHCI group ranged
between 5.8 and 32% in corrected conciliatory tendencies. The only comparable numbers for
chimpanzees come from the PREUSCHOFT et al. (2002) study and are generally higher (20 -
69.2%). However, the PREUSCHOFT et al. (2002) study contained eight immature individuals and
eight adults, including only one adult male. Additionally, only three of the eight adults partici-
pated in 20 or more conflicts (AM Jimoh: 31 conflicts; AF Mai: 25 conflicts; AF Anja: 28 con-
flicts). If we examine only these three adults the CCTs are 38.7, 20, and 50% respectively (the
PREUSCHOFT et al., 2002), giving a 36.2% mean corrected conciliatory tendency. It is very possi-
ble that the different composition of the groups at CHCI and Yerkes make direct comparisons of
the databases difficult.
In addition to individual corrected conciliatory tendencies we also examined dyads by
age/sex class. Female-female dyads had a substantially higher CCT than did male-male or het-
erosexual dyads. Again there is difficulty comparing these numbers to PREUSCHOFT et al. (2002)
due to the demographics of the two groups and the differential in adult-adult conflict pairs (262
at CHCI and 52 at Yerkes). However, female-female pairs had the highest corrected conciliatory
tendencies in both studies.
The inter-individual variation in reconciliation may be related to the relationships amongst
those individuals (ARNOLD & WHITEN, 2001 ; CORDS & AURELI, 2000; PREUSCHOFT et al., 2002).
At the CHCI there is variation across individual dyads in rates of reconciliation. For example,
the two lowest ranking individuals, who often interact affiliatively, had the highest dyadic CCT
(37.5%). This may reflect aspects of the "relationship quality" factor that is suggested to be
associated with an increased likelihood of dyadic reconciliation (ARNOLD & WHITEN, 2001;
PREUSCHOFT et al., 2002).
There is a difference across conflict Levels 1 and 2 in conciliatory tendencies. This may be
due to the differential intensity (no physical contact in Level 1 and physical contact in Level 2).
Alternatively, this differential may be an artifact of frequency, as Level 2 conflicts where nearly
twice as common as Level 1. While the current low number of Level 3 conflicts prevents a
232 A. FUENTES et al.
more effective assessment of the relationship between conflict intensity and reconciliation it is
interesting to note that SCHINO et al. (1998) found no significant differences between reconcilia-
tion related to the lowest ("threats") and the highest ("physical assault") conflict categories in a
group of Macaca fuscata.
REDIRECTION
Re-directed aggression occurred significantly more often during post-conflict periods than
matched control periods. The majority of redirection can be attributed to one individual (Loulis)
and 63% of all redirection was directed towards human caretakers (see MALONE et al., 2000, for
a detailed description).
CONSOLATION
Although affiliation occurred after conflicts, individuals did not engage in affiliative behav-
ior with third parties more so after conflicts than in matched control observations. However, it
may be relevant to note that 97 (48.9%) of the 198 potential consolation events in the PC and
MC were with human partners. The lack of significant difference in PC and MC consolation
mi ght suggest that for this group of chimpanzees consolation is not a common strategy for
negotiating conflict. This reduced role for consolation is supported by the free-ranging study at
Budongo (ARNOLO & WHITEN, 2001) where a lack of consolation after conflict was also
reported. We are currently exami ni ng the directionality of initiation of consolation behavi or
across all data collection periods in an effort to more accurately assess third part post-conflict
contact (i.e. DAS, 2000).
PROXIMITY
Conflict partners were spatially closer to one another sooner and more often during post-con-
flict observations than matched-control observations. This suggests an active manipulation of
space use following conflicts. One possible explanation for this result is that spatial proximity
and neutral behavior may be act as a peaceful post-conflict signal (SILK, 1996, 1998, 2000).
Therefore, remaining close and behaviorally neutral (not affiliative and not agonistic) may be
sufficient to clearly indicate the cessation of a conflict and the conflict participants' intent to not
engage in further conflict at the present time (DE WAAL & YOSHIHARA, 1983). ROWELL and
OLSON (1983) suggested that a change in spatial position communi cat ed a change in social rela-
tionship. Close proximity may allow a conflict partner to monitor their opponent s' movement s
and adjust their position accordingly. Alternatively, CORDS and AURELI (1993, 2000) reported
that maintaining close proximity after a conflict restored a functional aspect of a social relation-
ship.
VISUAL CONTACT
Visual contact between the conflict participants was significantly more common in the PC
than the MC period for this study. Overall this finding suggests that individuals may be actively
keeping the conflict partner in visual contact and monitoring his/her behavior. I f signals of
benign intent, or some alternative functional signal, are communi cat ed soon after a conflict it
would be very important for conflict participants to maintain visual contact.
Conflict and Post-conflict Behavior 233
POST-CONFLICT BEHAVIORS
At the group level the behaviors "threat" and "display" were exhibited significantly more
often after conflicts than during matched control observations across five of the six data collec-
tion sessions. This is due primarily to the behavioral contributions of Washoe and Loulis for
whom threat and display were common behaviors after conflicts. Interestingly, travel is a signif-
icant post-conflict behavi or at the group level for three of the four summer data collection ses-
sions but is absent at the group level during the 1999a and 2000a (winter) data collection
sessions when space was limited. This may indicate that individuals shift their pattern of behav-
ior as the available the space changes (JUDGE, 2000).
It is important to note that each individual exhibits specific behaviors significantly more
often in the PC periods across data collection periods and that three of the five individuals do
not appear to be consistent in these behaviors across the study (see Table 6). While this may be
a reflection of the sample size for each data collection period, it mi ght also reflect slight differ-
ences in group relationships, individual' s health, weather conditions, and other variables
impacting these chimpanzees.
CONCLUSI ON
This group of chimpanzees did reconcile approximately 17% of their conflicts. However,
individual variation in corrected conciliatory tendencies was large, suggesting that the mean
CCT may not accurately represent the strategies used by the individual chimpanzees in this
group. Individuals exhibited a range of behaviors, visually monitored former opponents, redi-
rected aggression and maintained increased proximity following conflicts. They did not appear
to seek or give consolation after conflicts. These results, combi ned with the relatively low cor-
rected conciliatory tendencies from the one other captive (BAKER & SMUTS, 1994) and one free-
ranging study (ARNOLD & WHITEN, 2001) suggest that post-conflict behavi or aside from
reconciliation may also be important to our understanding of chimpanzee conflict negotiation.
Given the variation in mean corrected conciliatory tendencies across the four published studies
(see Table 4) and the individual variation in CCT reported here and by PREUSCHOFT et al. (2002)
it is important to continue and expand on the investigations into chimpanzee behavi or following
conflicts. In light of increased evidence of relationship qualities on post-conflict behavi or pat-
terns and the modifiability of reconciliatory behavi or through social experience a broader
approach to conflict negotiation studies in chimpanzees is needed (ARNOLD & WHITEN, 2001;
CORDS & AURELI, 2000; DE WAAL & AURELI, 2000; PREUSCHOFT et al., 2002). Our data suggest
that proximity/use of space, visual monitoring, redirection of aggression, conflict pairs' rela-
tionships, and aspects of the captive environment may be important facets in the conflict negoti-
ation of this small group of chimpanzees.
Acknowledgements. We wish to thank Dr. ROGER FOUTS and DEBORAH FOUTS, Co-Directors of the
Chimpanzee and Human Communications Institute at Central Washington University, for their support and
encouragement. We thank all of the CHC| apprentices, students and staff who worked long and hard col-
lecting and entering the data, without them this project could not have taken place. We also wish to thank
Dr. F1LIPPO AURELI, Dr. ROBERT SUSSMAN, MARCEL HARVEY, and two anonymous reviewers for comments
and suggestions on various drafts of this manuscript. Finally, we wish to thank the chimpanzees of the
CHCI, Washoe, Loulis, Dar, Moja, and Tatu for making this project possible.
234 A. FUENTES et al.
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- - Received: July 24, 2001; Accepted: April 24, 2002
Authors' Names and Present Addresses: AGUSTIN FUENTES, Department o[Anthropolog); Notre Dame University, Notre
Dame, hMiana 46556-5639. U. S. A. e-mail: anthro@nd.edu; NICHOI.AS MAI.ONI-, Primate Behavior and Ecology
Program, Central Washington University, Ellensburg, Washington 98926-7544, U. S. A.; CRICKI;lq'I~ SANg, Department
c~l: Anthropolog); Washington Universit3; St. Louis, Missouri, U. S. A.; MEGAN MATHESON, Primate Behavior and
Ecology Program and Department o, f Psycholog3; Central WashhNton University Ellensburg, Washington 98926-7544.
U. S. A.; LORIt~N VAU(~HAN, San Diego WiM Animal Park, San Diego. Cal[/ornia, U. S. A.