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Carol Berman

Office: Spaulding Bldg 6, Room 165
Phone Number: (716) 645-0429
Email Address:


PhD, University of Cambridge
BA, Brandeis University

Research Interests

Primate social behavior, parent-offspring relationships, animal behavior, evolution of behavior, kinship, conflict management, ethological methods.

Courses Offered

Undergraduate Courses
APY 246 | Introduction to Primate Behavior
APY 344 | Animal Communication
APY 444 | Ethology Practicum  

Graduate Courses
APY 518 | Primate Social Behavior
APY 544 | Animal Communication
APY 547 | Ethology Practicum 
APY 550 | Evolution Colloquium

Current Research Projects

Parent-offspring Relationships and Social Development: Free-ranging rhesus monkeys on Cayo Santiago, Puerto Rico

Dr. Berman's long term research concerns parental behavior and infant social development. Berman is particularly interested in the complex inter-relationships between maternal behavior, early social relationships, social structure and demography. How do young animals become integated into the social structure of their group? How do social structure and demography influence maternal style and social development? What are the consequences of particular maternal styles for the infant’s developing social network and for the mother’s future reproductive performance? How are individual behavior patterns and social structure passed on from generation to generation?

Conflict Management, Kinship and Social Structure: Tibetan Macaques at Mt. Huangshan, China

Since 2000, Dr. Berman has focused on understanding the behavioral ecology and social structure of a relatively little studied species of macaque–wild Tibetan macaques in China. Working in collaboration with Consuel Ionica and Jinhua Li, Berman has investigated dominance style, post-conflict affiliation (reconciliation), patterns of male competition and cooperation, and variations in the expression of female kin bias in this species. We have also investigated patterns of conflict prevention, third party post conflict affiliation, and behavioral stress indicators among adults. A common aim in the studies is to test and refinement of theories concerned with the evolutionary origins of social diversity among human and nonhuman primates.

The Impact of Tourism on Primate Behavior and Conservation

Dr. Berman's study group of Tibetan macaques has been monitored by various researchers since 1986 and has been used as a tourist attraction since 1994. This have given Berman and her colleagues the opportunity to assess the effects of management for tourism by comparing behavior and infant mortality before tourist management began, during several years of management and during a temporary suspension of management in 2003. The group found markedly higher rates of infant mortality and aggression among adults in the provisioning area in years of tourist management. Many of the infants died from wounds following fights among the adults. The group has been examining specific aspects of tourist management to determine which of them may be responsible for increases in aggression and mortality.

Selected Publications

Please see Dr. Berman's bibliography for a reference of all publications.


B. Chapais and C. M. Berman.  KINSHIP AND BEHAVIOR IN PRIMATES. Oxford University Press, New York, 520 pp., 2004.

Peer-Reviewed Journal Articles and Book Chapters

K.N. Balasubramaniam , E.S. Dunayer, L.J. Gilhooly, K.A. Rosenfield and C.M. Berman. Group Size, Contest Competition, and Social Structure in Cayo Santiago Rhesus Macaques. BEHAVIOUR, 151: 1759-1798, 2014.

A. Yanagi and C.M. Berman. Functions of multiple play signals in free-ranging juvenile rhesus macaques (Macaca mulatta). BEHAVIOUR, 151: 1983-2014, 2014.

C.M. Berman. Primate kinship: contributions from Cayo Santiago. AMER. J. PRIMATOL. Featured article for a special edition, Online: DOI=10.1002/ajp.22383&ArticleID=2177526.

K.N Balasubramaniam, K, Dittmar, C.M. Berman, M., Butovskaya, M.A. Cooper, B. Majolo, H. Ogawa, G. Schino, B.Thierry, and Waal. Hierarchical Steepness and Phylogenetic Models I: Phylogenetic Signals in Macaca, ANIMAL BEHAVIOUR., 83: 1207-1218, 2012.

K.N Balasubramaniam, K, Dittmar, C.M. Berman, M., Butovskaya, M.A. Cooper,B. Majolo, H. Ogawa, G. Schino, B.Thierry, and Waal. Hierarchical Steepness, Counter-aggression and the Macaque Social Scale. AMERICAN J. PRIMATOLOGY, 74: 915-925, 2012.

 A. Yanagi and C.M. Berman. Gestural signals during social play in free-ranging rhesus macaques (Macaca mulatta): A systematic analysis. AMERICAN JOURNAL OF PRIMATOLOGY, Published online: DOI: 10.1002/ajp.22219, 9/30/13.

C.M. Berman and B. Thierry. Variation in Kin Bias: Species Differences and Time Constraints in Macaques, BEHAVIOUR, 147: 1863-1887, 2010.

 C. M. Berman. Primate Kin Preferences: Explaining Diversity. In: C. A. Salmon and T. K. Shackelford (Eds.), The Oxford Handbook of Evolutionary Family Psychology. New York: Oxford University Press, pp.248-278, 2011.

K.N. Balasubramaniam, C.M. Berman, H. Ogawa and J.H. Li. Using biological markets principles to examine patterns of grooming exchange in Macaca thibetana. AMERICAN J. PRIMATOLOGY, 73: 1269-1279, 2011.

C.M. Berman. Kinship: Family Ties and Social Behavior. In: Primates in Perspective, 2 edition. Edited by C.J. Campbell, A Fuentes, K.C. MacKinnon, M. Panger and S.K. Bearder. Oxford University Press, New York, pp. 576-587, 2011.

C.M. Berman, C.S. Ionica, M. Dorner and J-H Li. Post-conflict affiliation between former opponents in Macaca thibetana: For males only? INTERNATIONAL J. PRIMATOLOGY, 27: 827-854, 2006.

C.M. Berman, J-H Li, H. Ogawa, C.S. Ionica and H. Yin. Primate tourism and infant risk among Macaca thibetana at Mt. Huangshan, China. INTERNATIONAL J. PRIMATOLOGY, 28:1123-1141, 2007.

C.M. Berman, C.S. Ionica and J-H Li. Male tolerance and support among male Tibetan macaques (Macaca thibetana) on Mt Huangshan, China. BEHAVIOUR, 144: 631-661, 2007

C.M. Berman, C.S. Ionica and J-H Li. Dominance style among Tibetan macaques (Macaca thibetana) on Mt Huangshan, China. INTERNATIONAL J. PRIMATOLOGY, 25(6): 1283-1312, 2004.

 C. Chauvin and C. M. Berman. Intergenerational transmission of behavior. In: HOW SOCIETIES ARISE: THE MACAQUE MODEL, edited by B. Thierry, M. Singh and W. Kaumanns, Cambridge University Press, Cambridge, UK, pp. 209-234, 2004.

 C.M. Berman. Developmental Aspects of Kin Bias. In: KINSHIP AND BEHAVIOR IN PRIMATES. Edited by B. Chapais & C.M. Berman, Oxford Univ. Press, NY, pp. 317-346, 2004.

R.L. Johnson, I. Malik and C.M. Berman. On the quantification of suckling intensity in primates. AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY, 105: 33-42, 1998.

C.M. Berman and E. Kapsalis. Development of kin bias among rhesus monkeys: Maternal transmission or individual learning? ANIMAL BEHAVIOUR, 58: 883-894, 1999.

E. Kapsalis and C.M. Berman. Models of affiliative relationships among free-ranging rhesus monkeys (Macaca mulatta) I. Criteria for kinship. BEHAVIOUR, 133:1209-1234, 1996.

 E. Kapsalis and C.M. Berman. Models of affiliative relationships among free-ranging rhesus monkeys (Macaca mulatta) II. Testing predictions for three hypothesized organising principles. BEHAVIOUR, 133:1235-1263, 1996.

C.M. Berman, K.L.R. Rasmussen and S.J. Suomi. Group size, maternal behavior and social networks: A natural experiment with free-ranging monkeys. AN. BEHAV., 53:405-421, 1997.

C.M. Berman, K.L.R. Rasmussen and S.J. Suomi. Reproductive consequences of rejecting and non‑rejecting care patterns during estrus for free‑ranging rhesus monkey mothers.  BEHAV. ECOL. SOCIOBIOL., 32: 391‑399, 1993.

R.L. Johnson, C.M. Berman and I. Malik.  An integrative model of the lactational and environmental control of mating in female rhesus monkeys. ANIMAL BEHAV., 46:63‑78, 1993.

C. M. Berman.  Intergenerational transmission of maternal rejection rates among free‑ranging rhesus monkeys on Cayo Santiago. ANIMAL BEHAVIOUR, 39:329‑337, 1990.

C.M. Berman.  Maternal condition and offspring sex ratio in a group of free‑ranging rhesus monkeys: An eleven year study. AMERICAN NATURALIST, 131:307‑328, 1988.

C.M. Berman.  The ontogeny of social relationships between infant monkeys and group companions: I. Social networks and differentiation. ANIMAL BEHAV. 30:149‑162, 1982.