RESEARCH ARTICLE
Juvenile Social Relationships Reflect Adult Patterns of Behavior in Wild
Geladas
CAITLIN L. BARALE
1
*, DANIEL I. RUBENSTEIN
1
, AND JACINTA C. BEEHNER
2,3
1
Department of Ecology and Evolutionary Biology, Princeton University, Princeton, New Jersey
2
Department of Psychology, University of Michigan, Ann Arbor, Michigan
3
Department of Anthropology, University of Michigan, Ann Arbor, Michigan
Unlike many mammals, primates spend much of their lives as reproductively-immature juveniles.
During the juvenile period, they develop social relationships and physical skills that both facilitate
survival to adulthood and impact adult fitness. In this study, we use 2 years of observational data to
examine the development of these skills across the juvenile period in a wild cercopithecine primate,
the gelada (Theropithecus gelada). As adults, male and female geladas require different skills to be
successful; we therefore expected sex differences in social behavior and partner choice during the
juvenile period to already reflect these sex-specific trajectories. For example, males, who disperse at
puberty and ultimately must challenge other adult males for access to mates, should invest in high-
energy play-fighting with other males to develop fighting and rival assessment skills. In contrast,
philopatric females, who remain with their close kin throughout their lives, should invest more in
forming less-physical and more-social bonds with other females within their group. As predicted, sex
differences that foreshadowed sex-specific adult roles were apparent in play rates, the average number
of play partners per individual, grooming partner types and social partner preferences. Males played
more and had more play partners than same-age females. Males also groomed more often with
individuals from outside their natal group than females, although no sex difference was observed in
either grooming rates or number of grooming partners per individual. Females stopped playing earlier
than males, and instead invested in grooming relationships with close relatives. Additionally, we found
that individual play and grooming rates were temporally consistent for both males and females (i.e.,
from one year to the next year), suggesting that individuals exhibit stable behavioral phenotypes. We
conclude by discussing how early life in geladas may shape adult behavior and reproductive strategies.
Am. J. Primatol. 77:1086–1096, 2015. © 2015 Wiley Periodicals, Inc.
Key words: juvenile social behavior; partner preference; play; sex-specific behavior; male
dispersal
INTRODUCTION
A lengthy juvenile period distinguishes primates
from other mammals. Young primates spend the
time between weaning and sexual maturation
developing species-specific and sex-specific behav-
iors, building relationships, and skills for adult life,
and learning to negotiate their complex social and
physical environments [Bekoff, 1984; Byers, 1998;
Martin and Caro, 1985; Thompson, 1998]. Despite
the demonstrated importance of this developmental
period, primate research typically focuses on the
adults of a species. Juveniles are notoriously
challenging to study in the wild: their small
body size makes them difficult to spot and individu-
ally identify, they move unpredictably, and they are
constantly growing and changing. Yet, research on a
diverse set of non-primate taxa has linked juvenile
behavior and sociality to fitness-related skills such
as territory establishment, successful reproduction,
Contract grant sponsor: Wildlife Conservation Society;
contract grant sponsor: National Science Foundation;
contract grant numbers: BCS-0715179, BCS-1154314, IOS-
1255974; contract grant sponsor: Leakey Foundation;
contract grant sponsor: American Society of Primatologists;
contract grant sponsor: International Society of Primatologists;
contract grant sponsor: National Geographic Society;
contract grant numbers: Gr.# 8100-06, 8989-11;
contract grant sponsor: Princeton University;
contract grant sponsor: University of Michigan
Correspondence to: Caitlin L. Barale, 106A Guyot Hall,
Princeton University, Princeton NJ. E-mail: cbarale@prince-
ton.edu
Received 2 March 2015; revision accepted 9 June 2015
DOI: 10.1002/ajp.22443
Published online 26 June 2015 in Wiley Online Library
(wileyonlinelibrary.com).
American Journal of Primatology 77:1086–1096 (2015)
© 2015 Wiley Periodicals, Inc.