PRIMATES,17(3): 291-300, July 1976

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A Comparative Analysis of Subgroup Structure and Spatial Relationships in Captive Baboons and Squirrel Monkeys
LYNN FAIRBANKS

University of Washington
ABSTRACT. Subgroup structure, spacing patterns, and the relationship between spatial and behavioral interations were compared for captive baboons and squirrel monkeys. Analysis of the structure, intensity, and permanence of subgroups revealed that the baboons formed low intensity, overlapping subgroups which were relatively flexible while the squirrel monkeys segregated into permanent, high intensity, mutually exclusive cliques. Clique associations and social proximity relationships were found to be better predictors than dominance rank of the nature and frequency of behavioral interactions in the two colonies. INTRODUCTION Differences between primate species in typical spacing patterns and subgrouping tendencies have been noted since the early days of primatology. To mention just a few of these variations, hamadryas baboon (Papio hamadryas) troops subdivide into harems containing one male and several females (KUMMER, 1968); gibbons (Hylobates lar) typically live in family groups with one male, one female, and their young (CARPENTER, 1940); and rhesus macaques (Macaca mulatta) are usually found in multi-male troops with subdivisions along matrilineal kinship lines (ShoE, 1965). Variations have also been noted within species studied in different habitats, with langurs (Presbytis entellus) being found in one-male, multi-male, and all male troops (JAY, 1965; SUGIYAMA, 1964). While these differences in socio-spatial relationships have long been recognized as an important dimension in understanding primate social organization, there has been surprisingly little quantitative data on details of subgroup structure and the relationship of social spacing to behavioral processes. Most of the information presently available on primate spatial patterns comes from field studies and is primarily qualitative in nature with a few notable exceptions. KCMMZR(1968) collected data on both spatial distance and social interactions to precisely define the one-male subunits of the hamadryas baboon. SADE (1972) adapted several sociometric techniques to identify grooming networks and their relationship to dominance rank in free-ranging rhesus macaques on Cayo Santiago. These methods of quantitatively evaluating subgrouping patterns have only recently been applied to laboratory studies of primate behavior (FAIRBANI(S, 1974a). Complex analysis of spatial relationships in captive primate groups can not only be useful in describing species-typical social organization (MASON, 1971; ROSENBLUM, KAUFMAN, & Sa'VNES, 1964), but can also be used to monitor behavioral processes occurring within the group. Spatial structure of a captive group is both a consequence and a cause of social interactions, a consequence in that subgrouping patterns may

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be determined by past affiliative or hostile interactions between individuals, and a cause in that nonrandom spatial distribution influences the probability of future behavioral events. Differences between species in spacing patterns and their concommitant social behaviors can provide insight into the relationships and interactions between different styles of social and spatial behavior. The present study was designed to describe socio-spatial relationships in two primate species, savanna baboons (Papio cynocephalus) and squirrel monkeys (Saimiri sciureus). Prior information on spatial patterns in these two species in the field indicates that both live in multi-male groups of comparable size range, but that the subgroup structure of the two species is quite different. Baboons tend to aggregate in relatively large groups with no distinct substructure throughout most of their range (HALL & DEVORE, 1965; ROWELL, 1966a) while squirrel monkey troops have been reported to divide into smaller units, usually containing individuals of the same agesex class, to forage or socialize (TnoRINGTON, 1968; BALDWIN, 1971). TWO methods of analysis have been chosen to describe and compare spacing patterns and social relationships of these two species under captive conditions. First, a Clique analysis was designed to identify subgroups in a matrix of sitting associations, to assess the intensity and stability of these subgroups, and to evaluate the behavioral processes responsible for maintaining them. And second, a Social Proximity measure was developed to relate individual differences in the amount of time spent in social proximity to frequency and quality of behavioral interactions and dominancesubordinate relationships. PROCEDURE
SUBJECTS

Subjects were one social group of savanna baboons (Papio cynocephalus) containing eight adult females and one adult male, and one group of squirrel monkeys (Saimiri sciureus) with six adult females and two adult males. In addition to the adults observed as focal subjects, the baboon colony contained one juvenile female, one juvenile male, one brown infant, and four black infants. Two additional infants were born into the group during the course of the present study. The squirrel monkey colony contained one juvenile male and one infant at the onset of the study, with another infant being born during the course of data collection. All adults of both species were feral born and all infants and juveniles were born in captivity.
HOUSING

The baboon colony was maintained at the University of Washington Primate Field Station at Medical Lake, Washington. The group was housed in a large room, 11' 22' 8', with benches along the walls and pipes in the ceiling for sitting perches. Observations were made through a plexiglass window from an adjacent room. The squirrel monkey colony was maintained at the University of Washington Psychology Department in a wire-mesh cage, 8' 6' 4', with several branches and dowels for sitting perches. This group was observed from an adjacent room through a one-way glass.

Subgroup Structure and Spatial Relationships in Baboons and Squirrel Monkeys

OBSERVATION SCHEDULE

293

Data from the squirrel monkey colony were taken during 45 days of observation between September 1 and December 1, 1972. During an observation day, continuous data were recorded for each adult member of the colony, in random order, for 5 minutes. Data on the baboon colony were collected for 45 days between January 15 and April 15, 1973, according to the same observation schedule.
CODING SYSTEM

During a 5-minute observation period, data on spacing and behavioral interactions were recorded on a data sheet in sixty 5-second intervals. Spatial relationships were coded as: Solitary: No other animals within arm's reach. Near: Within arm's reach of another individual. Huddle: In gross body contact with another individual. In addition to a continuous record of spatial relationships, the following social behaviors were recorded when they occurred: Affiliative behaviors: including touch, lip smack, social groom, present, and affiliative behavior directed toward infants by non-mothers. Sexual behaviors: including genital inspection, mounting, and copulation. Agonistic behaviors: including threat, fear grimace, manipulation, and physical attack. Dominance behaviors: including displacement (a sequence where one individual leaves as the result of the approach of another individual) and genital display (abducting the thigh and exposing the genital region to another individual).
DATA ANALYSIS

In order to identify subgrouping patterns within each of the two colonies, the spacing data were analyzed by a method developed by FESTINGER(1949) and LUCE (1950) and used by SADE(1972) to identify grooming cliques in rhesus monkeys. A matrix was constructed of the percent of the total observation intervals that each adult in the group spent near or huddled with each other adult. This matrix was then analyzed to identify subgroups, or cliques, which included individuals who each reciprocally spent more than a specified amount of time near or huddled with each other individual in the clique. The intensity of clique relationships is determined by setting a discriminative value (DV), below which interactions are set at 0 and above which they are considered to be 1. (In SADE'S(Ioc. cir.) analysis the minimum number allowed in a clique was three individuals, but in the present study cliques of two members were also considered.) In addition to the subgroup analysis, the Social Proximity score was computed for each individual as a second measure of spatial association in the two colonies. Social Proximity was defined as the percent of the total timed intervals that each individual was recorded near or huddled with at least one other adult colony member. In order to relate spatial interactions with dominance-subordinate relationships, a measure of dominance was defined for each of the two species. The dominance rank

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for baboons was constructed according to the ability of each individual to displace each other individual. This method of ranking was found by ROWELL (1966b) to provide the clearest index of rank in a similar captive group of baboons. The squirrel monkeys were ranked according to the relative frequency of giving vs. receiving genital displays between individuals, a method used by several investigators (PLOOG, BLITZ, & PLOO6, 1963; CANDLANDet al., 1970) as an index of rank in this species. RESULTS CLIQUE STRUCTURE Figure 1 shows the subgroup structure for the baboon and squirrel monkey colonies. A clique analysis with a discriminative value (D V) of 10 % of the total time spent together in the baboon colony revealed four overlapping subgroups with two females remaining solitary. A similar analysis in the squirrel monkey colony detected
B A B O O N S S Q U I R R E L M O N K E Y S

B?

F?

2c~

DV = 25%

N OCLIQUES

@
l

%
DV: 50% N OCLIOUES %

Fig. 1. Clique structure of the baboon and squirrel monkey colonies at three levels of intensity.

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Table 1. Instances per individual per hour of affiliative, agonistic, and sexual behavior within and between subgroups for the baboon and squirrel monkey colonies.
Affiliative Baboons Within subgroups Between subgroups Squirrel monkeys Within subgroups Between subgroups ** p < . 0 1 5.49 3.14 ** 10.17 0.49 ** Agonistic 0.94 0.90 Sexual 0.50 0.50

0.17 0.07

0.50 0.22

two mutually exclusive cliques with one solitary male. Increasing the D V to 25 ~ to determine the intensity of clique associations revealed no high intensity cliques in the baboon colony. In the squirrel monkey colony the association of the adult male with all but one of the females in his clique dropped out at D V = 2 5 ~ , but the two adult female cliques remained intact. Both of these cliques were centered around a core of two females who each spent more than 50 ~ of their time together.
BEHAVIOR WITHIN AND BETWEEN SUBGROUPS

Table 1 summarizes behavioral interactions within and between subgroups (D V = 10~) for the two species. In the baboon colony there was almost twice as much affiliative behavior between individuals in the same subgroup as between individuals in different subgroups (t=3.43, p<.01). This effect was even more extreme in the squirrel monkey colony where 20 times as much affiliative behavior occurred within cliques as between (t=4.35, p<.01). In both species there was no detectable difference in the frequency of agonistic and sexual interactions by subgroup membership. Approximately half of the affiliative behavior within subgroups for both baboons and squirrel monkeys was focused around young infants. In the baboon colony, several changes of subgroup membership were noted following the birth of an infant. The amount of time ~ I spent near other females increased markedly post partum and her subgroup associations changed from only one to include four other females ( D V = 1 0 ~ ) . When s infant died two weeks later, her subgroup associations reverted immediately back to one. Following the birth of her infant, ~ E strengthened her existing clique associations and formed a new subgroup with the adult male. The two baboon females who were not included in any subgroups were the only two females in the group who did not have dependent infants at some point during the course of the study. While a considerable amount of affiliative attention was paid to infants in the squirrel monkey colony, the birth of an infant did not alter subgroup membership. The same females associated together with equal intensity before and after the births of their infants; females without infants confined their "aunting" behavior to members of their own subgroups.

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MALE N FEMALE ~ ' ~

BABOONS

O10

1020

2030

3040

4050

5060

6070

7080

8090

90 100

SQUIRREL M O N K E Y S

3
2

,N
0102030-

|
30 40
40so 5060 70 BO90-

1o

20

60

70

80

90

~0o

Fig. 2. Frequency histogram of Social Proximity scores for baboons and squirrel monkeys.

(Social Proximity score is the percent of 5 second intervals that each individual spent near or huddled with at least one other adult group member.) SOOAL PROXIMITY SCORE A frequency distribution of the Social Proximity scores (percent of timed intervals spent near or huddled with at least one other individual) is shown in Figure 2. Adult females in the baboon colony varied along a continuum on this measure, from 29 to 80 ~ . In contrast, the squirrel monkey females were uniformly high in the percent of time spent in Social Proximity with a range from 62 ~ to 86 ~ . In both species, males were relatively low in the amount of time spent near other group members.
DOMINANCE RANK

Analysis of relative displacements yielded an almost perfect linear dominance hierarchy in the baboon colony (Fig. 3). All dyads followed the most rigid definition of a dominance hierarchy where each individual was observed to displace each lower individual and be displaced by each higher individual in the dominance ranking, with one exception. No instances of displacements of ~ ~ by ~ c or ~ r) were recorded during the observation sessions, so the direction of other agonistic behaviors was used to determine ~ E'S rank. N o systematic ranking of the squirrel monkeys was possible based on direction of

Subgroup Structure and Spatial Relationships in Baboons and Squirrel Monkeys

O'~A

297

?,1,(z ? E ?,t.~ J ?+v ~" ~,vG ~4,n ? ~ Fig. 3. Dominance hierarchy of baboon group as determined by direction of spatial displacements.

genital displays. Only one genital display was recorded between the two adult males, and this was by the apparently less assertive male toward the more assertive. Between adult females, genital displays were relatively frequent in one subgroup but were reciprocal with no clear-cut linear ordering. No genital displays were recorded between females of the second subgroup.
BEHAVIORAL INTERACTIONS, SOCIAL PROXIMITY RANK, AND DOMINANCE RANK

The correlation between Social Proximity rank, Dominance rank, and frequency of affiliative and agonistic interactions for the baboon colony is shown in Table 2. There was a relatively high positive correlation between Social Proximity rank and involvement in both affiliative and agonistic behavior. In contrast, there was no linear relationship between Dominance rank and either behavioral category. A similar analysis was not performed for the squirrel monkey data as the Social Proximity scores were relatively invariant, especially among adult females, and a Dominance rank could not be reliably determined for the reasons explained above. Table 2. Spearman's rank order correlation between Social Proximity rank, Dominance rank, and affiliative and agonistic behavior in the baboon colony.
Social Proximity rank Dominance rank Affiliative behavior rho=.68 Agonistic behavior rho=.59

rho =.00

rho-- --.07

DISCUSSION Analysis of the clique associations of the captive baboon and squirrel monkey groups studied here revealed differences between these two species in subgroup intensity, stability, and structure. Adults in the captive baboon colony were organized in a series of low intensity, overlapping associations with no distinct subunits. One individual would frequently associate with several others sequentially, rather than simultaneously, so that most members were linked to most others indirectly through mutual sitting companions. Patterns of sitting together tended to be continuously graded so that the subgroups revealed by a clique analysis were relatively artificial breaks in a continuum rather than representing true structural units.

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The subgroup relations in the squirrel monkey colony were distinctively different from those found for the baboons. The squirrel monkeys segregated into clear-cut, mutually exclusive cliques with a high degree of association within and little or no interaction between units. Each of the two squirrel monkey subgroups contained a high intensity core of two adult females who spent more than 50 ~ of their total recorded time in close proximity to one another, with a third adult female who spent more than one-third of her time in close association with the other two. In addition to these structural features, the baboon and squirrel monkey subgroups also differed in relative permanence. Baboon subgroups were relatively flexible and were particularly influenced by the birth of an infant. In several cases in the baboon colony new subgroup relationships were formed when an adult female bore an infant and gradually broke up as the infant grew older or was removed from the group. In contrast, the squirrel monkey subgroups appeared to be relatively permanent and impervious to change. The subgroups described here have been constant for at least three years in spite of seasonal changes, births, deaths, and experimental manipulation following the termination of the observations reported here (FAIRBANKS,1974a, b). Data from the present study indicated that the loosely structured baboon subgroups and the more intense, exclusive squirrel monkey cliques were maintained primarily by affiliative behavior within the group. In the baboon colony, there was twice as much affiliative behavior directed toward other subgroups members as to outsiders, while in the squirrel monkey colony there was 20 times as much affiliative behavior within subgroups as between. Behavior between subgroups was characterized by lack of interaction rather than by any specific behaviors in both species. In the baboon colony, aggression was observed within subgroups as frequently as between, and there were no organized attacks of subgroup members against outsiders. With descriptive data of this nature, it is impossible to determine whether the spatial separation of non-interacting individuals was due simply to lack of positive attraction or to active avoidance based on past history of agonistic encounters. Experimental manipulation of spatial relationships would be necessary to determine the causal mechanisms for subgroup segregation in this species. For the squirrel monkeys, experimental analysis of subgroup processes in this same colony following the present study (FAIRBANKS,1974a) revealed that segregation of the females into separate groups was maintained by mutual avoidance rather than overt antagonistic behavior and that non-member males were excluded from female subgroups by infrequent, but intense, instances of aggression. Social Proximity analysis revealed differences between baboons and squirrel monkeys in the integration of each individual into the group spatial pattern. In the baboons there were marked individual differences in the amount of time spent in social proximity, with some females spending practically all of their time near other group members, and others remaining solitary. In contrast, squirrel monkey females were uniformly high in social proximity with only one of the males spending most of his time alone. An analysis of the relationship between Social Proximity rank and behavior

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produced remarkably high correlations within the baboon colony. The individuals who were the most social from a spatial standpoint were also involved in the most behavioral interactions, both positive and negative. The failure to find a relationship between dominance rank and the frequency of agonistic behavior is not as surprising as it may seem at first glance. Dominance rank is based on dyadic relationships and not on behavioral frequencies. Studies of this and other species (ROWELL, 1966b; KAUFMANN, 1967) have often noted that mid-ranking animals are more aggressive on the basis of frequency of aggression, but not in terms of the number of other individuals that they can dominate. While dominance rank may be a useful measure in understanding some features of baboon social organization, in the present situation Social Proximity rank proved to be a better predictor of day-to-day activity and events in the colony. In the squirrel monkey colony, an analysis of the direction of genital displays was not a useful measure of group structure and failed to produce a linear ranking between individuals. The signal function of the genital display is complex and it is only partially used as an assertion of rank (PLooa, 1967). In the present situation, displays between adult females were typically observed when one member returned to her habitual huddle group after a brief absence and appeared to function as a greetingrecognition ceremony between individuals. Subgroup structure proved to be the most useful indicator of the quality and direction of interactions for squirrel monkeys, with 88 % of all recorded behaviors between adults occurring within subgroup boundaries. Subgroup structure and measures like the Social Proximity rank used in the present study may provide a useful framework for organizing information on primate social systems. Dimensions such as intensity, composition, flexibility, and permanence of subgroups are relatively easily measured and compared across species, and can serve as sensitive measures of individual integration into a group. The present study has demonstrated the close tie between subgroup structure, social proximity, and the social dynamics of a captive group, and the usefulness of spacing concepts in predicting specific details of intra-group interactions in two primate species.

Acknowledgement. This work was supported by a National Science Foundation Graduate Fellowship.

REFERENCES

BALDWIN,J. D., 1971. The social organization of a semifree-ranging troop of squirrel monkeys (Saimiri sciureus). Folia primat., 14: 23-50. CANDLAND, D. K., D. C. BRYAN, K. KOPF, B. L. NAZAR & M. M. SENDOR, 1970. Squirrel moi~key heart-rate change during formation of status orders parallels the function in chickens. Jour. Comp. Physiol. Psychol., 70: 417-427. CARPENTER, C. R., 1940. A field study in Siam of the behavior and social relations of the gibbon (Hylobates lar). Comparat. Psychol. Monogr., 16:212 pp. FAIRBANKS, L., 1974a. An analysis of subgroup structure and process in a captive squirrel monkey (Saimiri sciureus) colony. Folia primat., 21: 209-224. - - , 1974b. Changes in the social role of a female with an infant in three primate

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species, Papio cynocephalus, Macaca nemestrina, and Saimiri sciureus, unpublished Ph. D. dissertation. FESTIN6ER, L., 1949. The analysis of sociograms using matrix algebra. Human Relations, 2: 153-158. HALL, K. R. L. & I. DEVoRE, 1965. Baboon social behavior. In: Primate Behavior: Field Studies of Monkeys and Apes, I. DEVoRE (ed.), Holt Rinehart & Winston, New York. JAY, P., 1965. The common langur of North India. In: Primate Behavior: Field Studies of Monkeys and Apes, I. DEVoRE (ed.), Holt Rinehart & Winston, New York. KAt~FMANN,J. H., 1967. Social relations of adult males in a free-ranging band of rhesus monkeys. In: Social Communication among Primates, S. A. ALTMANN (ed.), University of Chicago Press, Chicago. KUMMER, H., 1968. Social Organization of Hamadryas Baboons, A Field Study. University of Chicago Press, Chicago. LucE, R. D., 1950. Connectivity and generalized cliques in sociometric group structure. Psychometrika, 15 : 169-190. MASON W., 1971. Field and laboratory studies of social organization in Saimiri and Cal[ieebus. In: Primate Behavior: Developments in Field and Laboratory Research, L. A. ROSENBLUM (ed.). Vol. 2. Academic Press, New York. PLOOG, D. W. 1967. The Behavior of squirrel monkeys as revealed by sociometry, bioacoustics, and brain stimulation. In: Social Communication among Primates= S. A. ALTMAnN (ed.), University of Chicago Press, Chicago. ~ , J. BLITZ, & F. PLOOG, 1963. Studies on social and sexual behavior of the squirrel monkey (Saimiri sciureus). Folia Primat., 1 : 29-66. ROWELL, T. E., 1966a. Forest living baboons in Uganda. Jour. ZooI. Soe. Lond., 149: 344364. ~ , 1966b. Hierarchy in the organization of a captive baboon group. Anim. Behav., 14: 430-443. SADE, D. S., 1965. Some aspects of parent-offspring and sibling relations in a group of rhesus monkeys, with a discussion of grooming. Amer. Jour. Phys. Anthrop., 23: 1-17. ~ , 1972. Sociometrics of Maeaca mulatta. I. Linkages and cliques in grooming matrices. Folia Primat., 18 : 196-224. SU6IVAMA, Y., 1964. Group composition, population density, and some sociological observations of Hanuman langurs (Presbytis entellus). Primates, 5 : 7-38. THORIN~TON, R. W., 1968. Observations of squirrel monkeys in a Columbian forest. In: The Squirrel Monkey, L. A. ROSENBLUM& R. W. COOVER (eds.), Academic Press, New York. - - R e c e i v e d March 1, 1975; Accepted May 24, 1975 Author's Address: LYNN FAmBANKS,Department of Psychiatry, University of Cal(fornia at Los Angeles, Los Angeles, California 90024, U.S.A.