Professor; Director of the Primate Behavior Laboratory
Chair, Behavioral and Brain Sciences Program
604 Psychology Building
(706) 542-3036

EthoCebus Field Project


Ph.D. University California-Davis, 1978

Research Interests

Flexible instrumental behavior (problem-solving), manipulation, foraging and feeding in primates. I consider the developmental bases of flexible behavior, and the contribution of social learning and social context to the expression of flexible behavior. I work primarily with capuchins (Cebus; South American monkeys). I collaborate with many colleagues studying wild tool-using capuchin monkeys in Brazil, and in laboratory studies of spatial cognition, perception and action.

Selected Publications


Fragaszy, D.M., Visalberghi, E., & Fedigan, L. (2004). The Complete Capuchin. Cambridge University Press.

Fragaszy, D., & Perry, S. (Eds). (2003). The Biology of Traditions: Models and Evidence. Cambridge University Press.

Refereed Publications (from 2000 - present) Please note that these articles are for personal use only


Adams-Curtis, L.E., Fragaszy, D.M., & England, N. 2000. Prehension in infant capuchins (Cebus apella) from six weeks to twenty-four weeks: Video analysis of form and symmetry. American Journal of Primatology, 52, 55-60.

Christel, M., & Fragaszy, D. 2000. Manual function in (Cebus apella). Digital mobility, preshaping, and endurance in repetitive grasping. International Journal of Primatology 21, 697-719.

Cooper, M., Bernstein, I., Fragaszy, D., and de Waal, F. 2000. Integration of new males into four social groups of tufted capuchins (Cebus apella). International Journal of Primatology, 226, 663-683.

Dettmer, E., and Fragaszy, D. 2000. Determining the value of social companionship to captive tufted capuchin monkeys (Cebus apella). Journal of Applied Animal Welfare Science, 3, 293-304.

Fragaszy, D., & Visalberghi, E. 2000. Recognizing a swan: Socially-biased learning. Psychologia, 44, 82-98.


Johnson-Pynn, J., & Fragaszy, D. 2001. Do apes and monkeys rely upon conceptual reversibility? A review of studies using seriated nesting cups in children and nonhuman primates.Animal Cognition, 4, 315-324.


Fragaszy, D., Galloway, A., Johnson-Pynn, J., Hirsh, E., & Brakke, K. 2002. The sources of skill in seriating cups in children, monkeys, and apes. Developmental Science, 5, 118-131.


Fragaszy, D., Johnson-Pynn, J., Hirsh, E., & Brakke, K. 2003. Strategic navigation of two-dimensional alley mazes: Comparing capuchin monkeys and chimpanzees. Animal Cognition, 6, 149-160.

Johnson-Pynn, J., Fragaszy, D.M., & Cummins-Sebree, S. 2003. Common territories in comparative and developmental psychology: The quest for shared means and meaning in behavioral investigations. International Journal of Comparative Psychology, 16, 1-27.

Leighty, K., & Fragaszy, D. 2003. Joystick acquisition in tufted capuchin (Cebus apella) Animal Cognition, 6, 141-148.

Leighty, K., & Fragaszy, D. 2003. Primates in cyberspace: Using interactive computer tasks to study perception and action in nonhuman animals. Animal Cognition, 6, 137-139.

Fragaszy, D. 2003. Making space for traditions. Evolutionary Anthropology, 12, 61-70.


Fragaszy, D.M., & Visalberghi, E. 2004. Socially biased learning in monkeys. Learning and Behavior, 32, 1, 24-35.

Leighty, K.A., Byrne, G., Fragaszy, D.M., Visalberghi, E., Welker, C., & Lussier, I. 2004. Twinning in tufted capuchins (Cebus apella): Rate, survivorship, and physical development. Folia Primatologica, 75, 14-18.

Fragaszy, D.M., Izar, P., Visalberghi, E., Ottoni, E.B., & Gomes De Oliveira, M. 2004. Wild capuchin monkeys (Cebuslibidinosus) use anvils and stone pounding tools. American Journal of Primatology, 64, 359-366.


Rosengart, C. R., and Fragaszy, D.M. 2005. Experience and materials affect combinatorial construction in tufted capuchins monkeys. Journal of Comparative Psychology, 119, 166-178.

alloway, A., Addessi, E., Fragaszy, D., and Visalberghi, E. 2005. Social facilitation of eating familiar food in tufted capuchin monkeys (Cebusapella): Does it involve behavioral coordination? International Journal of Primatology, 26, 181-189.

Cummins-Sebree, S., and Fragaszy, D. 2005. Choosing and using tools: Capuchins use a different metric than tamarins. Journal of Comparative Psychology, 119, 210-219.

Takeshita, H., Fragaszy, D., Mizuno, Y., Matsuzawa, T.,Tomonaga, M., and Tanaka, M. 2005. Exploring by doing. How young chimpanzees discover surfaces through action with objects.Infant Behavior and Development, 28, 316-328

Matheson, M.D., Fragaszy, D.M., Johnson-Pynn, J.S. 2005. Response to novel housing in two groups of captive tufted capuchin monkeys (Cebus apella). Primates, 46, 235-240.

Fragaszy, D., and Cummins-Sebree, S. 2005. Relational spatial reasoning by a nonhuman: The example of capuchin monkeys.Behavioral Cognitive Neuroscience Reviews, 4, 282-306.


Izar, P.,Verderane M.P., Visalberghi E. , Ottoni E.B. , Gomes De Oliveira M, Shirley J ; Fragaszy D. 2006. Cross-genus adoption of a marmoset (Callithrix jacchus) by wild capuchin monkeys (Cebus libidinosus): case report. American Journal of Primatology, 68, 692-700.


Visalberghi, E., D. Fragaszy, E. Ottoni, P. Izar, M. G. de Oliveira, F. R. D. Andrade. 2007. Characteristics of hammer stones and anvils used by wild bearded capuchin monkeys (Cebus libidinosus) to crack open palm nuts. American Journal of Physical Anthropology, 132(3), 426-444.

Brakke, K., Fragaszy, D, Simpson, K., Hoy, E., and Cummins-Sebree, S. 2007. The production of unimanual and bimanual percussion in 12- to 24-month-old children. Infant Behavior and Development 30, 2-15.

Fragaszy, D. 2007. Relational spatial reasoning and tool use in capuchin monkeys. A primatologia no Brasil 10, 521-546.


Gunst, N., Boinski, S., and Fragaszy, D. 2008. Acquisition of foraging competence in wild brown capuchins (Cebus apella), with special reference to conspecifics' foraging artefacts as an indirect social influence. Behaviour 45(2), 195-229.

Hoy Kennedy, E., and Fragaszy, D. 2008. Analogical reasoning in a capuchin monkey. Journal of Comparative Psychology. 122(2), 167-175

Leighty, K., Menzel, C.,and Fragaszy, D. 2008. How young children and chimpanzees (Pan troglodytes) perceive objects in a 2D display: Putting an assumption to the test. Developmental Science. 11(5), 778-792

Visalberghi, E, Sabbatini, G, Spagnoletti, N, Andrade, FRD, Ottoni, E, Izar, P, Fragaszy, D. 2008. Physical properties of palm fruits processed with tools by wild bearded capuchins (Cebus libidinosus). American Journal of Primatology 70(9), 884-891.

Resende, B., Ottoni, E.B., and Fragaszy, D. 2008. Ontogeny of manipulative behavior and nut-cracking in young capuchin monkeys (Cebus apella): A Perception-action perspective.Developmental Science 11(6), 828-840.


Liu, Q., Simpson, K., Izar, P., Ottoni, E., Visalberghi, E., and Fragaszy, D. 2009. Kinematics and energetics of nut-cracking in wild capuchin monkeys (Cebus libidinosus) in Piaui, Brazil.American Journal of Physical Anthropology 138(2), 210-220.

Visalberghi, E., Addessi, E., Truppa, V., Spagnoletti, N., Ottoni, E., Izar, P., and Fragaszy, D. 2009. Selection of effective stone tools by wild bearded capuchin monkeys. Current Biology 19, 213-217.

Visalberghi, E., Spagnoletti, N., Ramos da Silva, E. D., Andrade, F. R. D., Ottoni, E., Izar, P., Fragaszy, D. (2009). Distribution of potential suitable hammers and transport of hammer tools and nuts by wild capuchin monkeys. Primates, 50, 95-104.

Crast, J., Fragaszy, D., Hayashi, M., and Matsuzawa, T. 2009. Dynamic in-hand movements in adult and young juvenile chimpanzees (Pan troglodytes). American Journal of Physical Anthropology 138(3), 274-285.

Fragaszy, D., Kennedy, E., Murnane, A., Menzel, C., Brewer, G., Johnson-Pynn, J., Hopkins, W. 2009. Navigating two-dimensional mazes: Chimpanzees (Pan troglodytes) and capuchins (Cebus apella sp.) profit from experience differently. Animal Cognition12, 491-504.

Lucas, P., Constantino, P., Chalk, J., Ziscovici, C., Wright, B., Fragaszy, D., Hill, D., Lee, J., Chai, H., Darvell, B., Lee, Pl, Yuen, T. (2009). Fallback foods: field assessment of mechanical properties and their potential effect on the dentition. American Journal of Physical Anthropology, 114, 643-352.

Wright, B. W., Wright, K. A., Chalk, J., Verderane, M. P., Dragaszy, D., Visalberghi, E., Izar, P., Ottoni, E. B., Constantino, P., & Vinyard, C. (2009). Fallback Foraging as a Way of Life: Using Dietary Toughness to Compare the Fallback Signal Among Capuchins and Implications For Interpreting Morphological Variation. American Journal of Physical Anthropology, 140, 687-699.


Gunst, N.Boinski, B., & Fragaszy, D. (2010). Development of skilled detection and extraction of embedded preys by wild brown capuchin monkeys (Cebus Apella). Journal of Comparative Psychology, 124, 194-204.

Gunst, N., Leca, J-B., Boinski, S., & Fragaszy, D. (2010). The Ontogeny of Handling Hard-to-Process Food in Wild Brown Capuchins (Cebus apella apella): Evidence From Foraging on the Fruit of Maximiliana maripa. American Journal of Primatology, 72, 960-973.

Fragaszy, D., Pickering, T., Liu, Q., Izar, P., Ottoni, E., and Visalberghi, E.  2010.  Bearded capuchin monkeys’ and a human’s efficiency at cracking palm nuts with stone tools: Field experiments.  Animal Behaviour, 79, 321-332.

Fragaszy, D. M., Greenberg, R., Visalberghi, E., Ottoni, E. B., Izar, P., & Liu, Q. (2010). How wild bearded capuchin monkeys select stones and nuts to minimize the number of strikes per nut cracked. Animal Behaviour, 80, 205-214

Crast, J., Hardy, J., & Fragaszy, D. 2010.  Inducing traditions in captive capuchin monkeys (Cebus apella).  Animal Behaviour, 80, 955-964.


Simpson, E., Varga, K., Frick, J., & Fragaszy, D. 2011.  Infants experience perceptual narrowing for nonprimate faces.  Infancy, 16(3), 318–328.  doi: 10.1111/j.15327078.

Liu, Q., Fragaszy, D., Wright, B., Wright, K., Izar, P. & Visalberghi E. (2011).  Wild bearded capuchin monkeys (Cebus libidinosus) place nuts in anvils selectively.  Animal Behaviour 81, 297-305.

Pan, J., Kennedy, E.H., Pickering, T., Menzel, C., Stone, B.W., & Fragaszy, D.M. 2011.  Development of maze navigation by tufted capuchins (Cebus apella).  Behavioural Processes,  86, 206–215. Doi:10.1016/j.beproc.2010.11.006

Fragaszy, D., & Simpson, E. (2011). Understanding emotions in primates: In honor of Darwin's 200th birthday.  American Journal of Primatology, 73, 1-4. doi:10.1002/ajp.20933 

Colbert- White, E.N., Covington, M.A., & Fragaszy, D.M. (2011). Social context influences the vocalizations of a home-raised African Grey parrot (Psittacus erithacus erithacus).  Journal of Comparative Psychology, 125(2), 175-184. doi:10.1037/a0022097

Fragaszy, D., Deputte, B., Cooper, E., Colbert-White, E., & Hemery, C. (2011).  When and how well can human-socialized capuchins match actions demonstrated by a familiar human? American Journal of Primatology, 73 (7), 643–654.  DOI: 10.1002/ajp.20941

Souto, A., Bione, C.B.C., Bastso, M., Bezerra, B., Fragaszy, D., & Schiel, N. (2011). Critically endangered blonde capuchins fish for termites and use new techniques to accomplish the task.Biology Letters,7(4), 532-535. 

Spagnoletti, N., Visalberghi, E., Ottoni, E., Izar, P., & Fragaszy, D. (2011). Stone tool use by adult wild bearded capuchin monkeys (Cebus libidinosus). Frequency, efficiency and tool selectivity.Journal of Human Evolution, 62(1), 97-107.   [clear] doi:10.1016/j.jhevol.2011.02.010 

Fragaszy, D., Stone, B., Scott, N., & Menzel, C. (2011).  How tufted capuchin monkeys (Cebus apella spp) and common chimpanzees (Pan troglodytes) align objects to surfaces: insights into spatial reasoning and implications for tool use.  American Journal of Primatology.   DOI: 10.1002/ajp.20966

Izar, P., Verderane, M., Peternelli-dos-Santos, L., Mendonca-Furtado, O., Presotto, A., Tokuda, M., Visalberghi, E., & Fragaszy, D. (2011). Flexible and Conservative Features of Social Systems in Tufted Capuchin Monkeys: Comparing the Socioecology of Sapajus libidinosus and Sapajus nigritus. American Journal of Primatology, 73, 1–17. DOI 10.1002/ajp.20968

Izar, P., Verderane, M., Peternelli-dos-Santos, L., Mendonça-Furtado, P., Presotto, A., Tokuda, M., Visalberghi, E., & Fragaszy, D. (2011). Flexible and conservative features of social systems in tufted capuchin moneys: comparing the socioecology of Sapajus libidinosus and Sapajus nigritus. American Journal of Primatology, 74, 315-331.


Massaro, L., Liu, Q., Visalberghi, E., & Fragaszy, D. (2012). Wild bearded capuchin (Sapajus libidinosus) select hammer tools on the basis of both stone mass and distance from the anvil. Animal Cognition. Doi: 10.1007/s10071-012-0530-x.

Spagnoletti, N., Visalberghi, E., Verderane, M.P., Ottoni, E.B., Izar, P., Fragaszy, D. 2012. Stone tool use in wild bearded capuchin monkeys (Cebus libidinosus). Is it a strategy to overcome food scarcity? Animal Behaviour, 83, 1285-1294.

Fragaszy, D. (2012). Community resources for learning: How capuchin monkeys construct technical traditions. Biological Theory. DOI: 10.1007/s13752-012-0032-8.

Duarte, M., Hanna, J., Liu, Q., Fragaszy, D. (2012). Kinematics of bipedal locomotion while carrying a load in the arms in beaded capuchin monkeys (Sapajus libidinosus). Journal of Human Evolution. DOI:10.1016/j.jhevol.2012.10.002


Fragaszy, D.M., Liu, Q., Wright, B.W., Allen, A., Brown, C.W. (2013). Wild bearded capuchin monkeys (Sapajus libidinosus)strategically place nuts in a stable position during nut-cracking. PLOS ONE, 8(2): E56182. DOI: 10, 1371/JOURNAL.PONE.0056182.

Verderane, M.P., Izar, P., Visalberghi, E., Fragaszy, D.M. (2013). Socioecology of wild bearded capuchin monkeys (Sapajus libidinosus): an analysis of social relationships among female primates that use tools in feeding. Behaviour, 150, 659-689.

Visalberghi, E., Haslam, M., Spagnoletti, N., Fragaszy, D. (2013). Use of stone hammer tools and anvils by bearded capuchin monkeys over time and space: Construction of an archeological record of tool use. Journal of Archeological Science, 40, 8, 3222-3232.

LaCour, L., Stone, B., Hopkins, W., Menzel, C., Fragaszy, D. (2013). What limits tool use in nonhuman primates? Insights from tufted capuchin monkeys (Sapajus spp.) and chimpanzees(Pan troglodytes) aligning three-dimensional objects to a surface. Animal Cognition, DOI 10.1007/s10071-013-0643-x.

Colbert-White, E.N., McCord, E.M., Sharpe, D.I., & Fragaszy, D.M. (2013). String-pulling behavior in a Harris’s Hawk Parabuteo unicinctus. The International Journal of Avian Science, doi: 10, 1111/ibi.12040.

Fragaszy, D., Biro, D., Eshchar, Y., Humle, T., Izar, P., Resende, B., and Visalberghi, E. 2013. The fourth dimension of tool use: temporally enduring artifacts aid primates learning to use tools. Philosophical Transactions of the Royal Society B, 368, 20120410.

Book Chapters 


Lacreuse, A., & Fragaszy, D.M. 2000. Exploration perceptive tactile chez les primate non humains. In Toucher pour connaitre. Psychologie cognitivede la perception tactile manuelle, eds. Hatwell, Y., Streri, A. and Gentaz, E. pp. 225-241. Presses Universitaires de France, Paris.


Visalberghi, E. & Fragaszy, D. 2002. Do monkeys ape? Ten Years After. In Imitation in Animals and Artifacts, eds. K. Dautenhahn & C. Nehaniv , pp. 473-499. MIT Press, Cambridge.


Fragaszy, D., & Perry, S. 2003. Towards a biology of traditions. In Traditions in Nonhuman Animals: Models and Evidence, eds. D. Fragaszy & S. Perry, pp. Cambridge University Press, Cambridge.


Cummins-Sebree, S.E., & Fragaszy 2005. Capuchins as stone-knappers?: an evaluation of the evidence. In Knapping Stone: A Uniquely Hominid Behavior, eds. V. Roux and B. Brill, pp. 171-182. Cambridge University Press, Cambridge.


Visalberghi, E. & Fragaszy, D. 2006. What is challenging about tool use? The capuchin's perspective. In Comparative Cognition:Experimental Explorations of Intelligence, eds. E. Wasserman & T. Zentall, pp. 529-552. Oxford University Press, Oxford.

Leighty, K.A., Fragaszy, D.M., & Brown, J.M. 2006. Darwin in Cyberspace: Advances in technology give us new avenues to study the continuity of cognition across species. In Primate Perspectives, ed. D. Washburn, pp. 37-46. American Psychological Association, Washington, D.C.


Fragaszy, D.M. & Crast, J. 2010. Monkeys and Prosimians: Social Learning. In: Breed, M.D. & Moore, J., (eds.) Encyclopedia of Animal Behavior, Vol. 2, pp. 468-474, Academic Press, Oxford.

Humle, T., & Fragaszy, D. 2010. Cognition and Tool Use. In: Campbell C, Fuentes A, MacKinnon K, Stumpf R, Bearder S. (eds.), Primates in Perspective, pp. 637-661. Oxford University Press, Oxford.


Visalberghi, E., & Fragaszy, D. 2011. Learning how to forage. Socially biased individual learning and “niche construction” in wild capuchin monkeys. In The Primate Mind, eds. F. de Waal and P.F. Ferrari, pp. 81-98. Harvard University Press, Cambridge, MA, pp. 81-98.


Visalberghi, E., & Fragaszy, D. 2012. What is challenging about tool use. The capuchin’s perspective. In Comparative Cognition: Experimental Explorations of Intelligence, 2nd Edition, eds. E. Wasserman and T. Zentall, Oxford University Press, pp. 777-799.

Fragaszy, D. & Liu, Q. 2012. Instrumental behavior and problem solving. In N.M. Seel (Ed.), Encyclopedia of the Sciences of Learning (Vol. 3, pp. 1579-1582). Springer: New York, NY.


Visalberghi, E. & Fragaszy, D.  2013.  The EthoCebus project.  Stone tool use by wild capuchin monkeys.  In Multidisciplinary Perspectives on the Cognition and Ecology of Tool Using Behaviors, eds. Sanz, C., Call, J. & Boesch, C.  Cambridge University Press, Cambridge, pp. 203-222.