Frans B. M. de Waal

C. H. Candler Professor of Primate Behavior

Office: 399 Psychology Building

Phone: 404-727-7898

Fax: 404-727-0372


Additional Contact Information

Mailing Address:

Department of Psychology

36 Eagle Row
Emory University

Atlanta, GA 30322

Additional Websites


Dr. de Waal received his Ph.D. in Biology and Zoology from Utrecht University, the Netherlands, in 1977. He completed his postdoctoral study of chimpanzees while associated with Utrecht University, in 1981, and moved the same year to the USA. He has been a National Academy of Sciences member since 2004, and a Royal Dutch Academy of Sciences member since 1993. Time featured him in 2007 as one of the World's One Hundred Most Influential People. He is also the Director of Living Links at the Yerkes National Primate Research Center.


  • PSYC 552: NAB Proseminar: Animal Behavior and Evolution
  • PSYC 325 (BIOL 325): Primate Social Psychology

Other Affiliations


My research on primate behavior focuses on social complexity in the widest possible sense, including alliance formation, reciprocal exchange, reconciliation following aggression, deceptive communication, and responses to environmentally induced stress. This research has a distinctly comparative character; it is being pursued with chimpanzees, bonobos, several macaque species, and capuchin monkeys.

The methodology requires group-living captive animals, preferably under naturalistic conditions such as found at major zoos and research institutions.

The four main topics of current interest are:

  • Cultural learning: Stimulus enhancement, imitation, and group specific habits in chimpanzees. A collaboration with Dr. Andrew Whiten at St. Andrews University focuses on the learning of artificial fruits and the spreading of knowledge in chimpanzees.
  • Behavioral economics: A series of experiments on reciprocal altruism and cooperation with capuchin monkeys has been conducted, most recently including reactions to reward division (e.g. inequity aversion). Similar studies on chimpanzees are underway in collaboration with Dr. Sarah Brosnan at GSU.
  • Empathy: Observational study of empathy responses in chimpanzees (e.g. consolation of distressed individuals) as well as computerized experiments in which chimpanzees chose from among emotional scenes presented on a screen. Similar research is being conducted with Asian elephants and bonobos.
  • Communication: Experiments on audience effects in food calls and observations of gestural communication in great apes, such as bonobos and chimpanzees.

Presently, my research unit includes 4 postdoctoral associates, 1 paid technician, 3 graduate students, and a variable number of undergraduates and temporary assistants.

We work at both the Yerkes Field Station (with its 2,000 primates in outdoor enclosures) and in the NSF-sponsored Capuchin Lab at the Yerkes Main Center (near Emory Campus). The research has been or is being supported by NIH, NSF, NIMH, and the Greater Good Center. We have collaborations with Zoo Atlanta, Bronx Zoo, Chester Zoo, St. Andrews University, Kyoto University, Chimp Haven (Louisiana), as well as other primate facilities, Thai elephant parks, and Lola ya Bonobo in the DRC.

Apart from being an active research unit, we also try to communicate the importance of primatological research to the general public, doing so both through the Living Links Center's website and via books for popular audiences (see Recent Publications).

Current Lab Members:

  • Dr. Darby Proctor, Postdoctoral Research Associate,
  • Dr. Victoria Horner, Postdoctoral Research Associate:
  • Dr. Matthew Campbell, Postdoctoral Research Associate,
  • Dr. Zanna Clay, Postdoctoral Research Associate,
  • Malini Suchak, graduate student, NAB,
  • Taylor Rubin, graduate student, NAB,
  • Sarah Calcutt, graduate student, NAB,
  • Julia Watzek, Research Associate,
  • Dr. Katie Hall, affiliate,

Many of my graduate students have supplemented their regular fellowships with NSF fellowships (4 students) or support from the Wenner-Gren Foundation, Leakey Foundation, Woodruff Foundation, Fulbright program, etcetera.

Recent Ex-Members:

  • Dr. Filippo Aureli, Reader in Animal Behaviour, John Moores University, UK.
  • Dr. Peter Verbeek, Professor of Psychology, Myazaki Int. College, Japan.
  • Dr. Lisa Parr, Ass. Research Professor, Emory's Ctr Behav. Neuroscience
  • Dr. Stephanie Preston, Assistant Professor, University of Michigan - Ann Arbor.
  • Dr. Jessica Flack, Co-Director, Wisconsin Institute for Discovery, Madison, WI
  • Dr. Sarah Brosnan, Assistant Professor, Georgia State University.
  • Dr. Amy S. Pollick, Director Government Affairs for the APS, Washington, DC.
  • Dr. Kristin E. Bonnie, Assistant Professor, Beloit College, Wisconson.
  • Dr. Teresa Romero, Visiting Scientist, Tokyo, Japan
  • Dr. Ann Weaver, Program Chair of Research, Argosy University Sarasota
  • Kristi Leimgruber, graduate student, Yale University
  • Dr. Marietta Dindo, Postdoctral Associate, George Washington University
  • Dr. Jennifer Pokorny, Postdoctoral Associate, Univ of California-Davis
  • Dr. Joshua Plotnik, Postdoctoral Associate, Univ of Cambridge, UK
  • Dr. Stephanie Preston, Associate Professor of Psychology, Univ Michigan


Books & Edited Volumes Since 2007

de Waal, F. B. M. (2009). THE AGE OF EMPATHY: Nature's Lessons for a Kinder Society. New York: Harmony Books.

de Waal, F. B. M., & Ferrari, P. F. (2012). THE PRIMATE MIND: Built to Connect With Other Minds. Cambridge, MA; Harvard University Press.

de Waal, Frans (2013). THE BONOBO AND THE ATHEIST: In Search of Humanism among the Primates. New York: Norton.

Articles in General Science Journals Since 2007

de Waal, F. B. M. (2012). The antiquity of empathy. Science 336: 874-876.

de Waal, F. B. M., & Ferrari, P. F. (2010). Towards a bottom-up perspective on animal and human cognition. Trends in Cognitive Sciences 14: 201-207.

de Waal, F. B. M., Leimgruber, K., & Greenberg, A. R. (2008). Giving is self-rewarding for monkeys. Proceedings of the National Academy of Sciences, USA 105: 13685-13689.

Flack, J. C., & de Waal, F. B. M. (2007). Context modulates signal meaning in primate communication. Proceedings National Academy of Sciences, USA 104: 1581-1586.

Horner, V., Carter, D. J., Suchak, M., & de Waal, F. B. M. (2011). Spontaneous prosocial choice by chimpanzees. Proceedings of the Academy of Sciences, USA 108: 13847–13851.

Plotnik, J. M., Lair, R. C., Suphachoksakun, W., & de Waal, F. B. M. (2011). Elephants know when they need a helping trunk in a cooperative task. Proceedings of the Academy of Sciences, USA 108: 5116–5121.

Pokorny, J., & de Waal, F. B. M. (2009). Monkeys recognize the faces of group mates in photographs. Proceedings of the National Academy of Sciences, USA 106: 21539-21543.

Pollick, A. S., & de Waal, F. B. M. (2007). Ape gestures and language evolution. Proceedings of the National Academy of Sciences, USA 104: 8184-8189.

Romero, M. T., Castellanos, M. A., & de Waal, F. B. M. (2010). Consolation as possible expression of sympathetic concern among chimpanzees. Proceedings of the National Academy of Sciences, USA 107: 12110-5.

van Wolkenten, M., Brosnan, S. F., de Waal, F. B. M. (2007). Inequity responses of monkeys modified by effort. Proceedings of the National Academy of Sciences 104:18854 –18859.

Whiten, A., Spiteri, A., Horner, V., Bonnie, K. E., Lambeth, S. P., Shapiro, S. J., de Waal, F. B. M. (2007). Transmission of multiple traditions within and between chimpanzee groups. Current Biology 17: 1038-1043.

Other Selected Articles and Book Chapters Since 2007
Balasubramaniam, K. N., Dittmar, K., Berman, C. M., Butovskaya, M., Cooper, M. A., Majolo, B., Ogawa, H., Schino, G., Thierry, B., & de Waal, F. B. M. (2012). Hierarchical steepness and phylogenetic models: Phylogenetic signals in Macaca. Animal Behaviour xx: xx-xx.

Bonnie, KE & de Waal FBM (2007). Copying without rewards: socially influenced foraging decisions among brown capuchin monkeys. Animal Cognition 10: 283-292.

Campbell, M. W., and de Waal, F. B. M. (2011). Ingroup-outgroup bias in contagious yawning by chimpanzees supports link to empathy. PLoS-ONE 5: e18283.

Campbell, M. W., Carter, J. D., Proctor, D., Eisenberg, M. L., & de Waal, F. B. M. (2009). Computer animations stimulate contagious yawning in chimpanzees. Proc. Royal Society London B. 276: 4255-4259.

de Waal, F. B. M. (2008). Putting the altruism back into altruism: The evolution of empathy. Annual Review of Psychology 59: 279-300.

de Waal, F. B. M. (2009). Darwin’s last laugh. Nature 460: 175.

de Waal, F. B. M. (2010). For Goodness' Sake. Review of "The Price of Altruism," New York Times Book Review (July 11).

de Waal, F. B. M. (2011). What is an animal emotion? The Year in Cognitive Neuroscience, Ann. N.Y. Acad. Sci. 1224: 191–206.

de Waal, F. B. M. (2012). Research chimpanzees may get a break. Plos Biology 10: e1001291.

de Waal, F. B. M., & Pokorny, J. (2008). Faces and behinds: Chimpanzee sex perception. Advanced Science Letters 1: 99-103.

de Waal, F. B. M., Boesch, C., Horner, V., & Whiten, A. (2008). Comparing social skills of children and apes. Science 319: 569.

Dindo, M., Thierry, B., de Waal, F. B. M., & Whiten, A. (2010). Conditional copying fidelity in capuchin monkeys (Cebus apella). Journal of Comparative Psychology 124: 29-37.

Dindo, M., Whiten, A., & de Waal, F. B. M. (2009). In-group conformity sustains different foraging traditions in capuchin monkeys (Cebus apella). PLoS ONE 4: e7858.

Hattori, Y., Leimgruber, K., Fujita, K., & de Waal, F. B. M. (2012). Food-related tolerance in capuchin monkeys (Cebus apella) varies with knowledge of the partner's previous food-consumption. Behaviour xx: xx-xx.

Horner, V., Proctor, D., Bonnie, K. E., Whiten, A., & de Waal, F. B. M. (2010). Prestige affects cultural learning in chimpanzees. PLoS-ONE 5: e10625.

Pokorny, J., Webb, C., & de Waal, F. B. M. (2011). An inversion effect modified by expertise in capuchin monkeys. Animal Cognition 14: 839-846.

Romero, M. T., & de Waal, F. B. M. (2010). Chimpanzee consolation: Third party identity as a window on possible function. Journal of Comparative Psychology 124: 278 –286.

Thierry, B., Aureli, F., Nunn, C. L., Petit, O., Abegg, C., & de Waal, F. B. M . (2007). A comparative study of conflict resolution in macaques: Insights into the nature of trait co-variation. Animal Behaviour 75: 847-860.