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Relationships in six groups of rhesus monkeys II. Dyads

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 5 2 5 6 5 -575 (1980)
Relationsh
PS
n Six Groups of Rhesus Monkeys
II. DYADS
J.P. HANBY
MRC Unit on the Development and Integration of Behaviour, University
Sub-Department of A n i m l Behauwur, Madingley, Cambridge CB3 8AA
KEY WORDS Social relationships, Primate behavior,
Interaction patterns, Centrality, Compatibility, Deference,
Attraction, Familiarity
Dyadic relationships are described for rhesus monkeys living in
ABSTRACT
six groups. The kinds, frequencies, directionality, and overall patterning of interactions are examined over a twelve-month period for adult males, adult females,
and young monkeys. Kinship, sex, age, reproductive conditions, time, events, and
other social relations all affected dyadic relationships. Certain patterns of interactions characterized different relationships, such as those termed “special,”“tense,”
“affiliative,” “conflicting.”A component approach is recommended for the understanding of relationships based on an appraisal of attraction, familiarity, and
deference between individuals. The importance of assessing relationships in the
group context is stressed.
A social group can be characterized as a network of dyadic relationships. These networks
reveal patterns, the analysis and comparison of
which elucidates aspects of group social structure and dynamics (Hanby, ’80). However, such
networks rest on the dyadic relationships
themselves, which also deserve examination.
This paper describes the patterns of interactions between pairs of rhesus monkeys living in
social groups.
The description and analysis of dyadic relationships is complex, involving consideration
of the kinds, frequencies, directionality, and
patterning of various interactions over time
(Hinde, ’76). Even so, dyads are really a
simplification of the entire group network, because all pairs affect and are affected by other
individuals and relationships. Relationships
are constantly changing toeresponding to
growth, aging, health, and reproductive condition of individuals, season, events, and other
factors.
The study of relationships in primates necessitates simplification. Some investigators have
focused on particular dyads living in a group
(e.g., mother-infant pairs, Hinde and
Spencer-Booth, ’671, while others have separated dyads from the group in order to study the
relationship (e.g., mother-infant pairs, Jensen
et al., ’67).Another way to simplify has been to
focus on one particular interaction (e.g., groom-
0002-948318015‘204-0565$02.30 0 1980 ALAN R. LISS. INC.
ing, Sade, ’72) in order to characterize the relationships between individuals. In this study
the relationships of adult males, adult females,
and immature monkeys were examined, all in
the group context, and a number of different
interactions were used to characterize the different relationships. It is but a beginning, and
certainly more should be done. Nevertheless,
even at this preliminary stage, some principles
of dyadic relationships can be derived and are
discussed.
METHODS
Housing
The monkeys lived in six different groups,
each caged separately. The cages had indooroutdoor portions separated by a closable swinging door. Cages were cleaned in the mornings
and food set out. The basic diet consisted of
commercial monkey biscuits and unhulled sunflower seeds and peanuts. Supplementary food
(e.g., fruit, vegetables) was given in the afternoons. Further details are given elsewhere
(Hanby, ’80).
Group compositions
The ages, sexes, numbers, and genetic relations are shown for each group over the twelve
month study period in Table 1.The numbers of
Received December 4, 1978; accepted September 24, 1979.
565
566
J.P. HANBY
TABLE 1. Group compositions (July 1972 through June 19731.
Adult male
Adult females
Young males
Young females
2 4 yrs
s 4 yrs
s3yz yrs
s3’h yrs
Min-max. monkeys
together a t one time
Maternal kin units
Young relatediunrelated
to adult male
Billy’s
(B’s)
Mariner’s
Mercury’s
Fergus’
Tim’s
Yuri’s
(MS)
(HS)
(Fsl
(T’sl
(Y’S)
1
4
1
1
4- 5
1
2
2
1
3- 4
2- 3
2
1
1- 2
2
1
3- 4
2- 3
2
1
3
2- 3
2- 3
9
8- 9
7-8
8- 9
8- 9
9- 10
2:3:2
212
2:3:3
411
2:2:3
310
7
21 1
3:2:2
312
53
610
individuals vary because of births (51, deaths
(31, removals (4), or introductions ( 3 ) . Each
group is named after its single adult male.
Adult monkeys were 4 years of age or older;
young monkeys 3% years of age or younger, a t
the beginning of the study.
The maternal kin units refer to two or more
individuals genetically related through the
mother (“uterine kin”). A kinship unit of 2:2:3
means that the group had three units; two
mother-young dyads and the other a mother
with two offspring. The male’s relatedness to
the young in his group is also shown. Four of the
six young monkeys that were not offspring of
the group male were born into the group. The
other two unrelated youngsters were introduced with their mothers when less than one
month old. Effectively, then, all young present
in the group had never known any other environment. All adult group members had been in
their groups for a minimum of nine months
prior to the start of this study; most had lived in
the groups for several years.
Data collection
Data were collected between the hours of
0900-1300. Each group was observed for a
minimum of six hours per month for twelve
months (usually two hours per day for three
consecutive days). Eight different observers
scored behaviors onto checksheets from which
two kinds of data were later summarized:
events and activity samples.
Events were scored whenever they occurred,
and included: display, present, mount, aggression, avoidanceigrin (grimace). Activity samples were made every 5th minute during an
observation period. An individual was scored as
being ‘hlone” if 60cm. or further from another
monkey. Individuals were scored as in ‘)roximity” if within arm’s reach or less than 60cm.
4
The percentage of point samples on which individuals were found in proximity is used as an
estimate of time spent with one another.
Within the category of proximity, monkeys
were noted as being in contact (passive),grooming, play, or together without interacting in one
of these ways. Brief episodes of agonistic or
sociosexual interactions were rarely caught on
the 5th minute and were analysed as events.
Definitions of these behaviors are given in a
previous report (Hanby, ’80).The data base for
the study reported here was 7206 5th minute
point samples and 35,185 minutes (586.4
hours).
RESULTS
The total number of dyads living in six different groups was 223 (B’s = 36, M’s = 36, H’s =
35, F s = 36, T s = 35, Y’s = 45). The relationships of adult males (49 dyads) and those of
adult females with one another (39 dyads) receive the most detailed attention. The relationships between the adult females and young
monkeys (96 dyads) and those between the
young monkeys themselves (39 dyads) are considered more briefly. They have been and will
be described more extensively in a number of
reports on this colony of monkeys (e.g., Hinde
and Spencer-Booth, ’67; Hinde and White, ’74;
Hinde and Proctor, ’77; McGinnis, ’75; White,
’77; Simpson and Howe, in prep.).
Adult male relationships
The adult male was a dominant, if not
dominating, individual in every group. His relationships were extensive, often one-sided (not
reciprocal), and usually had a large agonistic
component. Proximity relations are summarized in Table 2, and some directional interactions between the six adult males and the
members of their groups are shown in Figure 1.
DYADIC RELATIONSHIPS OF MONKEYS
TABLE 2. Proximity relations of adult males.*
~~
~
Adult male - young monkey
Adult male
Adult female Related Unrelated Male Female
Mean%
(S.D.)
Dyads
16.3
(11.2)
25
(
11.6
7.4)
18
9.2
(8.1)
6
(
10.3 11.6
6.8) ( 8.3)
12
12
*Scores are the mean percent (andstandard deviation, S.D.) of samples
in proximity.
On average, the six males spent about half
their time alone and half in proximity to others
(X% samples alone = 50.8, S.D. = 13.5, N = 6
males). When they were with others, the males
clearly tended to spend most of their time in the
company of adult females. Males spent less
time with immature monkeys, and it did not
appear to make any difference whether the
young monkey was related or unrelated, male
or female. An adult male’s time in proximity to
immatures depended largely on how much time
he spent with the mother of the youngster. In 20
of the 21 instances, the adult male spent more
time in proximity to the mother than he did to
the young monkey. When the young was in
proximity to the male, the mother tended to be
also. The one exception was an unrelated juvenile male who was in proximity to the adult
male on 23% of the samples, while his mother
spent but 18%.
Five adult males had one particular adult
female with whom they spent most oftheir time
in proximity. The sixth male spent about a n
equal percentage with two adult females (B
with V and T). In all these cases of “preferred”
proximity partners, the time in proximity was
between 20% and 47%.Mutuality is implied in
the definition of proximity, yet in most cases
involving the adult male and especially in
these cases of a “preferred” proximity partner,
it was often the female who initiated the proximity.
Directionality in some aspects of adult male
relationships are shown in Figure 1. In all six
groups, the adult male was most often the recipient of grooming, avoidance, presenting, and
(not shown here) play; the actor in aggression
and (not shown) mounting. This overall pattern
of male-other relationships can be called deferential; in general, agonistic, sociosexual, and
grooming interacitons were consistently oneway. Reciprocity did occur especially in grooming with adult females; presenting and play
with young monkeys.
567
A closer look at the patterning of interactions
between adult males and females revealed a
special balance of components in certain dyads.
The overall relationship could be called close or
positive and involved high levels of proximity
and grooming, low levels of avoidance, and
especially important, a low ratio of avoidance to
aggression, plus a high ratio of grooming to
presenting. Each of the six males had this special patterning with a t least one female for
most of the duration of the study. The three
dyads with the most consistent patterning
(B-V, M-G, F-H) typify the relationship: proximity scores highest for male-female dyads (>
20%), grooming also highest (5-7% of all point
samples), grooming to proximity ratio high
(20-22% of time in proximity involved in
groomiiig), avoid to aggression ratio low (less
than 2) and grooming to presenting highest in
the group. Four other dyads had comparable
scores for most of one year, but certain components changed.
Close or special relationships with the male
seemed to result from a n active effort on the
part of the female t o maintain proximity, tolerating his high levels of aggression and avoiding him less frequently than other females. The
importance of grooming in maintaining this
relationship was obvious, and grooming, aggression, and presenting were more often reciprocated in these pairs than others. Maintaining
the relationship was thus not solely due to the
female and her special pattern of deference;
there was some degree of attraction displayed
by the male as well.
Some females appeared to work especially
hard to maintain or establish a close relationship with the male. These relationships could
be called “tense” and involved changing but
lower levels of proximity and grooming, frequent presenting, and high ratios of grooming
to proximity (20- 50%) and very high avoidance
to aggression (5-10 times as many avoids and
grins to the male as aggressions received from
him). Such a pattern suggested that the female
was attracted to the male, and yet she was
afraid of him and not relaxed in his company.
Other male-female dyads could be characterized by their low levels of positive interactions. Not only were levels of proximity and
grooming low, but the ratio ofgrooming to proximity or presenting was also low, while ratios of
avoiding to aggression were moderate, and aggression often more frequent than in other
pairs. Such a pattern implied that the female
was not trying to maintain or establish a closer
relationship with the male, accepting a rather
568
J.P. HANBY
limited and somewhat “evasive” relationship.
Recently introduced females, females without
living kin in the group, and young females
tended to have these evasive relationships with
the male. Thus, while unfamiliarity and aversion were factors, so was the degree of integration of the female into the group as a whole.
Since one female in the group “monopolized’
the male’s time, and she and her offspring and
friends interacted with him the most, other
females’relationships with the adult male were
definitely affected. Nevertheless, all the adult
females in any group managed to become pregnant from time to time, so the ones with a limited relationship had a t least a sexual component, even though there were few other positive
aspects.
Adult female relationships
Relationships between adult females and
with young monkeys were more complicated
than those involving the adult male. The influence of a large number of variables (e.g.,reproductive state, number of kin present, group
composition changes) were reflected in the
changing patterns of interactions that make up
adult female relationships. The effects of some
specific kinds of changes within groups have
been shown for entire networks elsewhere
(Hanby, ’80), and some examples will be given
shortly. First, some generalizations about
adult female relationships can be made based
on an examination of the patterning of interactions over a year.
Adult females were found alone on only
about a third of the samples (X% = 32.7, S.D. =
12.3, N = 23); thus, they spent a large proportion of their time in proximity to one or more
individuals. The percentage of time they spent
with particular others depended on relatedness, as shown in Table 3. Adult females were
most pften found in proximity to their own offspring, regardless of sex or age. The next highest proximity levels were with other kin, again
regardless of sex or age. On average, adult
females were in proximity to the adult male
(16%)more often than they were to unrelated
adult females or immatures (11%).
The females which usually had the highest
percentages of time in proximity to the male
(and a special relationship with him) also occupied a special place in the group as a whole.
These females all had at least one other female
TABLE 3. Proximity relations of adult females
with other adult females or young monkeys
according to relatedness.*
Offspring
~~~
Adult femaleAdult female
Mean %
(S.D )
Dyads
~
Non kin
Other kin
~-~
~~
20.4
3.8)
5
(
12.8
8.3)
32
(
19.1
7.3)
7
(
9.1
6.2)
31
(
22.2
7.6)
5
(
35.5
6.4)
2
(
Adult femaleYoung males
Mean %
(S.D.)
Dyads
46.1
(16.2)
10
Adult femaleYoung
females
-~~
Mean %
(S.D.)
Dvads
43.1
(13.9)
12
(
~~~
10.8
6.5)
31
*Scores are the mean percent (and standard deviation S.D.)of samples
in proximity.
with whom they spent 20- 50% of the samples
in proximity. In addition to the male and this
preferred female proximity partner, these
females also had high levels of time in proximity to their own offspring and the offspring of
their “friend.” Thus, these females were members of clear sub-groups or clusters and are
called central or “key”females. Other aspects of
their relationships are mentioned below.
All the 39 pairings of adult females with one
another were sorted as to the percentage of
samples found in proximity in order to facilitate an examination of other aspects of their
relationships. These proximity groupings are
shown in Table 4.It was immediately apparent
that proximity alone was not an adequate basis
for describing relationships; further subdivisions were made according to relatedness
for the high proximity group and the degree of
aggression for the medium and low proximity
groups.
Five of the seven related adult female dyads
fell in the high proximity category, and the
patterning of their interactions typify what
might be called relaxed and positive. Compared
with the close or special relationships with
adult males, these filial female pairs had the
same amount of time in proximity, but lower
Fig. 1. Dyadic relationships of adult males. Triangles enclose a male’s initial, circles enclose a female’s. Older
monkeys are a t the top of each sociogram, and younger monkeys arranged according to age downwards. Links
represent the direction (arrows) and frequency of grooming (per 1000 point samples), aggression, avoidigrin, and
presenting (each per 10000 minutes). The number of lines per link indicate relative frequency (i.e., 1-10 = one
broken line, 11-20 = one solid line, 21-30 = two lines, and so on, until > 60, shown by a solid band).
569
DYADIC RELATIONSHIPS OF MONKEYS
Groom
Agg rQssion
-
-Avoid/g rin
PrQsQnt
(XperlOOOOmins)
Billy-
u
MarinQr
A 0
Y m
1
Dyadic Relationships of Adult Males
570
J.P.HANBY
TABLE 4 . Relationships between adult females.*
Type of dyad
High proximity ( 3 20%)
Filial
5 dyads
Friendly
6 dyads
Interactions
Aggression
w1000
minutes
Av_oid/grin
x/1000
minutes
Presenting
X/lOOo
minutes
.96
,981
.98
(1.00)
( ,881
2.18
(2.98)
2.35
(2.47)
2.43
(1.61)
(
.55
,741
(
.61
.77)
.56
.76)
Rcximity
XI100
samples
Grooming
x/100
samples
27.2
6.6)
1.46
(1.07)
(
25.8
4.3)
26.4
(
(
.68
Mean
11 dyads
(S.D.)
( 5.2)
1.88
(2.23)
1.72
(1.92)
1.77
(1.51)
Medium proximity U O - l 9 % )
Companionable
9 dyads
(
15.0
2.6)
1.33
(1.05)
(
.92
,771
1.69
(1.20)
(
13.4
2.7)
1.65
(1.16)
6.64
(2.30)
4.26
(1.70)
Mean
17 dyads
(S.D.)
(
14.3
2.6)
1.50
(1.05)
3.59
(3.38)
2.90
(1.93)
(
Low proximity (< 10%)
Unfriendly
5 dyads
(
3.8)
(
52
.61)
(
1.44
.77)
1.20
(1.46)
( .09)
(
5.9
3.0)
(
.22
.35)
6.58
(3.07)
3.37
(2.27)
(
.08
.20)
(
4.2
3.5)
(
.36
,491
3.79
(3.60)
2.33
(2.33)
(
.24
.38)
Conflicting
8 dyads
Antagonistic
6 dyads
Mean
11 dyads
(S.D.)
5.2
.04
* A total of 39 dyads are sorted into three groups on the basis of the percent of samples found in proximity. Further subdivisions are based on kinship
(for high proximity category only1 or agonistic components (medium and low proximity categories only). Scores are mean frequency (standard
deviations) of various interactions for specified sample or time units.
levels of grooming (and a lower grooming to
proximity ratio), plus less aggression and
presenting and much less avoidance. These elements are what suggest the more relaxed nature of the relationship between filial pairs.
“Friends,” who also spent considerable time
in proximity, had higher levels of grooming and
agonistic interactions, implying that one or
both individuals were working relatively harder to maintain these relationships than were
the related females. Friendly female dyads all
had lower levels of those elements suggesting
tension than did the females with special relationships with the male. Of the six dyads
termed “friends,”four involved a female with a
special relationship with the male. All of the
central females had filial or friendly relationships with other females. With one exception,
these Biliative (filial or friendly) relationships
involved fairly consistent patterns of deference,
i.e., one female received grooming, avoidance,
presenting, while being the aggressor and
mounter. The dominant in the pair was the
older or the one who also had the special relationship with the adult male.
Like affiliative pairs, but a t lower levels of
activities, were nine pairs that could be called
companions (this category included the other
two, more distantly related dyads; aunt-niece,
cousins). Eight dyads had similar levels of
grooming and proximity to the companionable
pairs but had very high levels of agonistic and
sociosexual interactions. Such a balance of
negative and positive elements, plus the fact
that these females’ relationships were constantly changing over time and events, implied
that they were in conflict. In several of the
dyads, deference patterns were inconsistent.
These unsettled relationships often seemed to
be competitive, specifically in relation to the
adult male. Some females with conflicting or
competitive relations with certain other adult
females also had “tense” relationships with the
male.
DYADIC RELATIONSHIPS OF MONKEYS
571
Agonistic interactions were also a frequent other aspects of Polly’s initial relationships
component in some other adult female pairs, were positive, for she was a member of a large
but these “antagonistic” dyads had very low kin cluster, her mother was the group’s central
levels of proximity, grooming, and sociosexual female, and Polly was deferred to by most other
behavior, implying a more one-sided and nega- individuals and relatively relaxed with the
tive relationship. The remaining adult female adult male. In contrast, Eve had no kin living in
dyads all had very low frequencies of any kind her home group and had conflicting relationof interactions, though the negative elements ships with two other females and a tense one
were predominant. These “unfriendly” pairs with the male. Nevertheless, she did manage to
just didn’t seem to have any attraction to one maintain a companionable relationship with
another and spent their time and activities the central female.
with others. These individuals were often
After they were moved, the two females conperipheral or isolated group members.
tinued to have different kinds of relationships,
While the individuals themselves produced though both deferred to others in their new
these relationships by the ways in which they groups, both received high and similar levels of
interacted over time, there were some factors aggression and avoided others t h e same
that clearly influenced them. The effects of amounts. Because they were new, both females
other relationships (e.g., with the male or other were subjected to much harassment by others,
kin) on any particular relationship has been but they responded in different ways. Polly acshown (see also Hanby, ’80). Reproductive tively worked at establishing positive relationcondition was another obvious factor affecting ships (by grooming, presenting, playing, inrelationships of adult females. Estrous cycles itiating, and remaining in proximity); she was
produced short-term changes involving tempo- especially attentive to the adult male. Eve, on
rarily increased proximity, consorting, and the other hand, evaded others, especially the
copulation with the adult male. Pregnancy had adult male; her relationships remained narrow
varied effects, some females showing little and negative until the end of the study.
changes in relationships and others much more
Relationships of young monkeys
dramatic ones. Births generally produced
changes in relationships; often proximity with
Young monkeys spent about t h e s a g e
others increased, particularly proximity t o amount of time alone as did adult females (X%
other mothers with newborns, while the fre- = 34.2, S.D. = 13.4,N = 26). Of the time they
quency of avoidance also increased (see Hanby, spent in proximity to others, most was with
’80, and Hinde and Proctor, ’77 for specific their mothers. Other proximity relations are
shown in Table 5. Siblings and other relatives
examples).
Aging also appeared to affect relationships, had a high proportion of time together while
younger females generally having more ag- orphans, with no relatives present, spent their
onistic interactions, while older females time in proximity to peers. There was a clear
showed lower levels of a number of activities tendency for young males to spend more time in
except proximity. Familiarity was also a factor. proximity to male peers than females did with
The most consistent, positive pairings were be- peers of either sex.
tween long term associates; kin, or unrelated.
Offspring of mothers who had special reAmong relationships with a predominance of lationships with the adult male also spent more
negative or inconsistent aspects were dyads time with him than did other immatures in the
containing a relatively recently introduced group, played with him, were defended by him,
female. It was obvious that females had to and generally interacted in more ways. Simiinteract in certain ways if a positive relation- larly, immatures tended to interact more positively with adult females with whom their
ship were to be established and maintained.
An illustration of the way different females mothers had affiliative or companionable reestablished new relationships compared to how lationships. Offspring of conflicting or competithey maintained earlier ones is shown in Fig- tive mothers often had elements of antagonism
ure 2. Only proximity relations are shown in in their relationships as well. In short, the rethe diagram, but they exemplify the differences lationships of adults, especially kin, strongly
between these two females, both of whom were affected the relationships of immatures.
young adults without offspring, colony born,
Immature monkeys had relationships with
and not previously moved.
adults that were rather restricted in quality,
Polly clearly had closer and more extensive involving some proximity, usually little groomproximity relations in both her original and ing, but more agonistic and sociosexual innew group than did her counterpart, Eve. Most teractions. On the other hand, relationships be-
J.P. HANBY
572
I
i
\
/K\
/N\
In NQW Groups (Xof 4 mcs )
In Original GroupscX of 3mos)
Links indicatQ ‘lo of samplQs found in proximity ( < 6 0 c m )
_ _ _ _
5 O/O ,
~
5 - 9 % , =I O - I 4 % ,
15-1g0/o,QtC
Proximity Relationships and Group Changes
Fig. 2. Proximity relations and group changes for two adult females. Adults represented by dark symbols,
younger by lighter; males by triangles, females by circles.
tween the immatures themselves involved a terns of deference. Young monkeys generally
wide variety of interactions at higher levels deferred to adults, especially to the male, but in
(cuddling, playing, fighting, and sociosexual one group where adult relationships were not
behaviors), though proximity and grooming very close or positive, the juveniles’ relationwere usually a t lower levels than between ships were notably more unsettled and less defadults. The patterning of interactions con- erential. Another aspect of juvenile relationstantly changed in most relationships between ships affecting or being affected by adults was
immatures, just as the relations with their particularly noticable in the case of daughters
mothers and siblings changed. The detailed de- of the central or key female in the group. These
scription of these relationships will not be juvenile and adolescent females teased and
given here, and the reader is referred to citings “baited’ adults, especially the adult male; inof papers a t the beginning ofthe results section. terfered with play between other immatures
The relationships of some juveniles and ado- and the adult male, and generally made a nuilescents deserves special mention. One of the sance of themselves. They could also be very
characteristics of the changing relationships of friendly to adults and welcoming to new or
young monkeys seemed to be inconsistent pat- reintroduced group members, but relationships
DYADIC RELATIONSHIPS OF MONKEYS
TABLE 5. Proximity among young monkeys.*
Siblings or
other kin
Mean%
(S.D.)
Dyads
27.6
(
8.1)
9
Orphans- Male- Female- Malepeers
male female female
16.3
23.7
(10.8) (13.8)
10
7
15.0
(
8.4)
7
16.0
(
8.7)
25
* Scoresare meanpercent (standard deviation) ofsamples inproximity.
were, again, inconsistent. These young females
were often involved in agonistic interactions,
displayed often, and frequently enlisted the
support of the adult male in fights within the
group or when aggression was directed outside
the group. In sum, immature relationships
could have important and often disruptive effects on other rleationships, while relations
among adults clearly affected those of young
monkeys as well.
DISCUSSION
The findings reported here supplement those
of a variety of studies on groups, dyads, or social
structure (e.g., Cheney, ’ 7 6 ; Koyama, ’ 6 7 ;
Kummer, ’75; Kurland, ’77;Loy and Loy, ’74;
Missakian, ’72; Rosenblum, ’71; Sade, ’65, ’72;
Seyfarth, ’76, ’77; Simonds, ’74). The importance of kinship, age, and sex in determining
relationships has been widely demonstrated
and discussed. In addition, this study stresses
the importance of examining relationships in
the group context by more than one measure.
Certain patterns of interactions between individuals emerged from this study which were
given names in an attempt to characterize
them, e.g., “special,”“tense,” “affiliative,”“conflicting,” “evasive.” For each dyad, the kinds,
frequencies, and directionality of interactions
were important, but the overall patterning was
necessary to describe any relationship. The
ratios of particular interactions were meaningful: the ratio of grooming to proximity may
reflect the degree of tension in one another’s
company; the ratio of presenting to grooming
and proximity may indicate the ease with
which one member of a dyad could attract the
attention or company of another; the ratio of
avoidance to aggression seemed to reflect how
willing or able one individual was to tolerate
hostility. Combinations of ratios with levels of
proximity were one way of quantifying the degree of effort involved in maintaining (or establishing) a relationship.
Using such patterns to evaluate adult relationships led to the conclusion that females
had to work relatively harder to maintain close
relationships with the male than they did with
573
one another. (For the same percentage of time
in proximity, females groomed more, presented
more, and avoided much more, even though
aggression was about the same or lower for
dyads involving the male). Another conclusion
was that pairs of related adult females had the
most relaxed relationships of all. The generality of these and other results needs to be
known, and specifically, whether or not the patterns or balances of elements are similar, useful, and meaningful.
Consistencies in patterns of interactions
added another dimension to the characterization of relationships. One of the most important
appeared to be consistent deference of one
member of a dyad to the other. Usually, this
meant that one individual avoided, presented,
did not show aggression towards, and groomed
more often than the partner. Deference was
primarily a matter of directionality, though in
cases of change, levels of aggression, avoidance,
presenting, and grooming might rise. Changes
in deference patterns often had a rise in aggression relative to avoiding (i.e.,fights), combined
with little change or even increases in proximity scores. Such a pattern involved competitiveness more than antagonism. Inconsistencies in deference patterns were associated with
changes in reproductive state in females, group
composition changes, and health and age (the
most inconsistent deference patterns were seen
in immatures’ relationships). The most consistent deference was seen in relationships with
the adult male, for all others tended to defer to
him. Filial and friendly pairs of adult females
also had stable patterns of deference. Clear and
consistent deference may be a requisite for the
maintainance of close, positive relationships.
The relationships of certain females were
especially interesting, for one in every group
managed to have a close relationship with the
adult male and a t least one other adult female
(plus a generally positive relationship with her
own and her friends offspring). This female
was called “central,” and she usually deferred
to the male, while other females deferred to her.
Although she could also be called a “dominant”
female, that would place the emphasis on the
agonistic components and obscure what seemed
to be equal or even more important elements in
her relationships. These females appeared to
expend consistent effort (e.g., grooming,
presenting) in maintaining close relationships,
and often were rather less aggressive and
avoided less often than other females (especially so in relationships with kin). Central
females seldom had relationships with low
levels of interactions of any sort.
574
J.P. HANBY
The central female had a key position in the
group as a whole, and her relationships affected
other relationships. The position was a n important one in the structure of these groups, and it
would be interesting to know if the same kinds
of relations occur in larger groups, especially
those with more adult males. How necessary
the male is to the “centrality” of the female and
the extent of his preference for or acceptance of
the particular female remains ambiguous in
this study, though when he was temporarily
absent from a group, at least one central female
retained her status. Why certain females competed for the position and others did not is another problem. Success in maintaining a central position for the females i n this study
seemed to depend on familiarity or residence,
maturity, and the presence of kin, plus a n active effort. Presumably, the position is a valuable one, direct benefits being less effort spent
in avoiding others (however, aggression was
not low and certainly was a cost). Indirect benefits came to the female’s offspring who had
more interactions and rather special relationships with the adult male, and were more often
deferred to by other immatures and even adult
females. The social opportunities and confidence engendered may have direct benefits to
the offspring as it matures, as was seen in the
case of a daughter of a central female when
moved to a new group.
In addition to examining relationships in
other primates and situations, it is hoped that a
terminology for relationships can be developed
and concepts clarified. Some ideas gained by
this study of relationships of monkeys in small
groups suggests that there are a t least three
major components to any positive relationship:
attraction, familiarity, and deference. These
factors may apply to other species and have
certain theoretical implications. The terms are
arbitrary but can be briefly defined here.
( a ) Attraction. This is a continuum of
qualities, ranging from the attraction between
living things, through attractions due to mutual need (e.g., co-operation in food-getting or
defence), attractions due to age (e.g., mothers/
offspring) and sex (reproductive males and
females), and finally, the attraction between
two personalities (regardless of age and sex).
f b ) Familiarity. The degree of acquaintance
between any two individuals is fundamental to
the formation or maintenance of a relationship.
Familiarity means interacting in a variety of
ways over some extended period of time so that
the dyad members can learn to assess and pre-
dict each other’s behaviour. This component of
a relationship requires perceptual abilities
(e.g., memory of attributes) as well as some
degree of proximity or active communication.
(c) Deference. This word has been chosen to
represent a rather fuzzy array of qualities such
a s dominance-subordinance, compromise and
co-ordination in activities, control of motivation, inhibition of actions, awareness of mood,
temperament, and intent. In any compatible
and enduring relationship, one member defers
to another, or both defer a t different times or in
different situations. This is necessary because
no two mobile, social organisms are in precisely
the same state at the same time. Clear patterns
of deference facilitate t h e formation and
maintenance of a relationship. Unclear patterns seem bound to cause or be the result of
conflict and could be called incompatibilities.
Given the three factors coarsely described
above, one can see that several different types
of relationships can result from different balances. For instance, friends would have mutual
attraction, familiarity, and deference; enemies
would have low attraction, some familiarity,
but inconsistent deference; colleagues have
familiarity of a certain sort only, clear deference but maybe little attraction. Concise names
for other combinations of these elements are
difficult to find.
How various kinds of relationships are established, and how the weighting for different factors varies for individuals and species, is a n
important field open for investigation. Kummer (’75)found in his studies of gelada baboons
that agonistic and sociosexual interactions occurred earlier in the sequence of bond formation than did grooming. This suggests that
clear deferential patterns follow attraction and
must be established before the more positive
and relaxed gestures can be used in a relationship.
A study of relationships between male chimpanzees (Bygott, ’74) suggests that clear deference is necessary between two aggressive
males if they are to associate together. However, it was even more interesting that the
closest “friends” (other than siblings) differed
considerably i n their agonistic dominance rank
within the group. Clear deference was also a
component of close relationships in these captive rhesus monkeys.
Familiarity must depend on time and opportunity. Just how the spacing and duration of
interactions affects the formation and maintenance of relationships is another practically
unexplored field. Attraction is also a very broad
DYADIC RELATIONSHIPS OF MONKEYS
term that deserves more definition. How attraction is expressed and how it is affected by
other aspects of the relationship (e.g., imcompatibilities) and other relationships deserves
study. Since, among social primates, a dyadic
relationship exists in the context of a group,
there will always be competitive, tense, or incompatible elements in the relationships. One
adult female may wish to be with another, but
that other one may be in consort with a n aggressive male or surrounded by a large family
to which her attention is directed. Demands by
the adult female’s own infant will further compete with her desire and ability to be with that
female. All relationships are t h u s a compromise and a result of different forces of attraction and demand, not only within t h e
group, but from the environment as well (e.g.,
the amount of time spent feeding will influence
the time available for building familiarity and
the opportunities for forming alliances, etc.).
Studying the formation and maintenance of
relations in small, captive groups allows a focus
on the patterning and essential qualities of
dyadic relationships. It is a relatively simple
and productive way to proceed to some understanding of relationships in general. However,
certainly, more can be done of an experimental
nature in captive colonies and more in the way
of describing relationships in free-ranging
groups. Relationships need to be studied over
long time periods too. It is hoped that this paper
has contributed some data for comparison and
discussion and may stimulate attention to relationships that are considered in their proper
context, the social network.
ACKNOWLEDGMENTS
I am immensely grateful to David Bygott for
consistent support, encouragement, and even
typing; to Les Barden for help on the diagrams;
to L. White, D. Cheney, R. Seyfarth, M.
Simpson, and M. Thorndahl for comments on
the early drafts; to A. Simpson for help with
data preparation, and to Robert Hinde for the
opportunity, execution, and final reporting of
this study. The work was funded by the Medical
Research Council of England.
575
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