We use a number of non-invasive techniques to study the behavior and minds of animals. Here, Dr. Krupenye does eye-tracking with Kanzi, a bonobo living at Ape Initiative - a science and education nonprofit.
Publications
All papers are available for personal or academic use upon request ([email protected])
Key Publications
45. Warren, E.P, Pilgrim, C. P, Aellen, M. P, Morford, J. P, Herrmann, E., Mann, R. P., Biro, D., & Krupenye, C. (in press). Chimpanzees spontaneously prepare for mutually-exclusive possibilities and collective context strengthens this behavior. Philosophical Transactions of the Royal Society B
44. Bastos, A. P. M.P., & Krupenye, C. (2025). Evidence for representation of pretend objects by Kanzi, a language trained bonobo. Science, 391 (6785), 583-586.
43. Townrow, L.G & Krupenye, C. (2025). Bonobos point more for ignorant than knowledgeable social partners. Proceedings of the National Academy of Sciences, 122(6), e2412450122.
42. Carvajal, L.G & Krupenye, C. (2025). Mental representation of the locations and identities of multiple hidden agents or objects by a bonobo. Proceedings of the Royal Society B, 292: 20250640.
41. Lewis, L.S., Wessling, E.G., Kano, F., Stevens, J.M.G., Call, J., & Krupenye, C. (2023). Bonobos and chimpanzees remember familiar conspecifics for decades. Proceedings of the National Academy of Sciences, 120(52): e2304903120. doi: 10.1073/pnas.2304903120.
40. Kano, F., Krupenye, C., Hirata, S., Tomonaga, M., & Call, J. (2019). Great apes use self- experience to anticipate an agent’s action in a false belief test. Proceedings of the National Academy of Sciences, 116(42), 20904-20909. doi: 10.1073/pnas.1910095116.
39. Krupenye, C., and Hare, B. (2018). Bonobos prefer individuals that hinder others over those that help. Current Biology. 28(2): 280-286. doi: 10.1016/j.cub.2017.11.061.
38. Krupenye, C.*, Kano, F.*, Hirata, S., Call, J., and Tomasello, M. (2016). Great apes anticipate that other individuals will act according to false beliefs. Science, 354(6308): 110-114. doi: 10.1126/science.aaf8110. -selected by Science as one of the top ten breakthroughs of 2016
Other Publications
37. Morford, J. P, Lewin, P., Mann, R. P., Krupenye, C., & Biro, D. (in press). Collective route memories emerge through differential forgetting of navigational information in homing pigeons. Scientific Reports
36. Bastos, A. P. M.P, Foster, G.G, Wood, P. M., & Krupenye, C. (in press). Do dogs rationally infer the causes of failed actions? PLoS One
35. Lewis, L.S.G, Kano, F., Stevens, J.M.G., DuBois, J.G.G, Call, J., & Krupenye, C. (in press). Do great apes know each other’s names? Probing great ape comprehension of social vocal labels. Animal Behavior and Cognition
34. Townrow, L.G & Krupenye, C. (2025). Reply to Tauzin: How to implement novelty in theory of mind tasks. Proceedings of the National Academy of Sciences, 122 (21) e2503973122.
33. Townrow, L.G & Krupenye, C. (2025). Reply to Royka and Santos: Representing ignorance, and the evolution of theory of mind. Proceedings of the National Academy of Sciences, 122 (20), e2506762122.
32. Krupenye, C., Carvajal, L.G, & Bastos, A. P. M.P (2025). Psychological mechanisms for individual recognition- and anonymous-societies in humans and other animals. Behavioral and Brain Sciences, 48, e65.
31. Bastos, A.P.M., Warren, E., & Krupenye, C. (2024). What evidence can validate a dog training method? Learning & Behavior
30. Krupenye, C. (2024). Divergence in bonobo and chimpanzee social life. Learning & Behavior
29. Westra, E., Fitzpatrick, S., Brosnan, S., Gruber, T., Hobaiter, C., Hopper, L., Kelly, D., Krupenye, C., Luncz, L., Theriault, J., & Andrews, K. (2024). In search of animal normativity: a framework for studying social norms in nonhuman animals. Biological Reviews, 10.1111/brv.13056.
28. Lewis, L.S., & Krupenye, C. (2022). Theory of mind in nonhuman primates. In B.L. Schwartz & M.J. Beran (Eds.) Primate Cognitive Studies. Cambridge University Press.
27. Farrar, B., Altschul, D., Fischer, J., Tennie, C., Ostojic, L., Krupenye, C., & Rodrigo, A. (2022). Replication and reproducibility in primate cognition research. In B.L. Schwartz & M.J. Beran (Eds.) Primate Cognitive Studies. Cambridge University Press.
26. Lewis, L.S., & Krupenye, C. (2022). Eye-tracking as a window into primate social cognition. American Journal of Primatology. doi: 10.1002/ajp.23393.
25. Kano, F., Furuichi, T., Hashimoto, C., Krupenye, C., Leinwand, J.G., Hopper, L.M., Martin, C.F., Otsuka, R., & Tajima, T. (2022). What is so unique about the human eye? Comparative image analysis on the external eye morphology of human and nonhuman great apes. Evolution and Human Behaviour. doi: 10.1016/j.evolhumbehav.2021.12.004
24. Durdevic, K, & Krupenye, C. (2021). Representing knowledge, belief, and everything in between: Representational complexity in humans and other apes. Behavioral and Brain Sciences, 44: e150. doi: 10.1017/S0140525X20001855.
23. Amodio, P., Farrar, B.G., Krupenye, C., Ostojić, L., & Clayton, N. (2021). Little evidence that Eurasian jays protect their caches by responding to cues about a conspecific’s desire and visual perspective. eLife, 10, e69647. doi: 10.7554/eLife.69647.
22. Feldblum, J., Krupenye, C., Bray, J., Pusey, A.E., & Gilby, I.C. (2021). Social bonds provide multiple pathways to reproductive success in wild male chimpanzees. iScience, 24, 102864.
21. Lewis, L.S., Kano, F., Stevens, J.M.G., DuBois, J.G., Call, J., & Krupenye, C. (2021). Bonobos and chimpanzees preferentially attend to familiar members of the dominant sex. Animal Behaviour, 177, 193-206.
20. Hopper, L. M., Gulli, R. A., Howard, L. M., Kano, F., Krupenye, C., Ryan, A., & Paukner, A. (2020). The application of non-invasive, restraint-free eye-tracking methods for use with nonhuman primates. Behavior Research Methods. doi: 10.3758/s13428-020-01465-6.
19. Krupenye, C. (2020). The evolution of mentalizing in humans and other primates. In M. Gilead & K. Ochsner (Eds.) The Neural Basis of Mentalizing: A Social-Cognitive and Affective Neuroscience Perspective. Springer Press.
18. Kano, F., Call, J., & Krupenye, C. (2020). Primates pass dynamically social anticipatory-looking false belief tests. Trends in Cognitive Sciences, doi: 10.1016/j.tics.2020.07.003.
17. Bowie, A., Krupenye, C., Mbonzo, P., Minesi, F., & Hare, B. (2020). Implicit measures help demonstrate the value of conservation education in the Democratic Republic of the Congo. Frontiers in Psychology. doi: 10.3389/fpsyg.2020.00386.
16. Krupenye, C. & Call, J. (2019). Theory of mind in animals: Current and future directions. WIREs Cognitive Science, e1503. read-only link
15. Krupenye, C., Tan, J., & Hare, B. (2018). Bonobos voluntarily hand food to others but not toys or tools. Proceedings of the Royal Society B. 285(1886): 20181536. doi: 10.1098/rspb.2018.1536.
14. Kano, F., Moore, R., Krupenye, C., Hirata, S., Tomonaga, M., & Call, J. (2018). Human ostensive signals do not enhance gaze-following in chimpanzees but do enhance object-oriented attention. Animal Cognition. 21(5): 715-728. doi: https://doi.org/10.1007/s10071-018-1205-z.
13. Krupenye, C., MacLean, E. L., and Hare, B. (2017). Does the bonobo have a (chimpanzee-like) theory of mind? In B. Hare & S. Yamamoto (Eds.) Bonobos: Unique in Mind, Brain, and Behavior. Oxford University Press. DOI: 10.1093/oso/9780198728511.003.0006.
12. Kano, F.*, Krupenye, C.*, Hirata, S., Call, J., and Tomasello, M. (2017). Submentalizing cannot explain belief-based action anticipation in apes. Trends in Cognitive Sciences. 21(9): 633-634. doi: http://dx.doi.org/10.1016/j.tics.2017.06.011.
11. Krupenye, C.*, Kano, F.*, Hirata, S., Call, J., and Tomasello, M. (2017). A test of the submentalizing hypothesis: Apes’ performance in a false belief task inanimate control. Communicative & Integrative Biology. 10(4): e1343771.
10. Krupenye, C. (2017). Guesser-knower paradigm. Encylopedia of Animal Cognition and Behavior. doi: 10.1007/978-3-319-47829-6_1605-1.
9. Kano, F., Krupenye, C., Hirata, S., and Call, J. (2017). Eye tracking uncovered great apes' ability to anticipate that other individuals will act according to false beliefs. Communicative & Integrative Biology. 10(2): e1299836.
8. Reddy, R. B., Krupenye, C., MacLean, E. L., and Hare, B. (2016). No evidence for contagious yawning in lemurs. Animal Cognition, 19(5): 889-898. doi: 10.1007/s10071-016-0986-1.
7. Krupenye, C., Rosati, A. G., and Hare, B. (2016). What’s in a frame? Response to Kanngiesser & Woike (2016). Biology Letters, 12(1): 20150959. doi: 10.1098/rsbl.2015.0959.
6. Foerster, S., McLellan, K., Schroepfer-Walker, K., Murray, C., Krupenye, C., Gilby, I.C., and Pusey, A. (2015). Social bonds in the dispersing sex: partner preferences among adult female chimpanzees. Animal Behaviour, 105: 139-152. doi: 10.1016/j.anbehav.2015.04.012.
5. Krupenye, C., Rosati, A.G., and Hare, B. (2015). Bonobos and chimpanzees exhibit human-like framing effects. Biology Letters, 11(2). doi: 10.1098/rsbl.2014.0527.
4. MacLean, E., Krupenye, C., and Hare, B. (2014). Dogs account for body orientation but not visual barriers when responding to pointing gestures. Journal of Comparative Psychology, 128(3): 285-297. doi: 10.1037/a0035742.
3. Bray, J., Krupenye, C., and Hare, B. (2014). Ring-tailed lemurs (Lemur catta) exploit information about what others can see but not what they can hear. Animal Cognition, 17(3): 735-744. doi: 10.1007/s10071-013-0705-0
2. Rosati, A.G., Herrmann, E., Kaminski, J., Krupenye, C., Melis, A.P., Schroepfer, K., Tan, J., Warneken, F., Wobber, V. & Hare, B. (2012). Assessing the psychological health of captive and wild apes: A response to Ferdowsian et al. (2011). Journal of Comparative Psychology, 127(3): 329-336. doi: 10.1037/a0029144.
1. Gruber, T., Potts, K.B., Krupenye, C., Byrne, M., Mackworth-Young, C., McGrew, W.C., Reynolds, V., & Zuberbuhler, K. (2012). The influence of ecology on chimpanzee cultural behavior: A case study of five Ugandan chimpanzee communities. Journal of Comparative Psychology, 126(4): 446-57. doi: 10.1037/a0028702.
Key Publications
45. Warren, E.P, Pilgrim, C. P, Aellen, M. P, Morford, J. P, Herrmann, E., Mann, R. P., Biro, D., & Krupenye, C. (in press). Chimpanzees spontaneously prepare for mutually-exclusive possibilities and collective context strengthens this behavior. Philosophical Transactions of the Royal Society B
44. Bastos, A. P. M.P., & Krupenye, C. (2025). Evidence for representation of pretend objects by Kanzi, a language trained bonobo. Science, 391 (6785), 583-586.
43. Townrow, L.G & Krupenye, C. (2025). Bonobos point more for ignorant than knowledgeable social partners. Proceedings of the National Academy of Sciences, 122(6), e2412450122.
42. Carvajal, L.G & Krupenye, C. (2025). Mental representation of the locations and identities of multiple hidden agents or objects by a bonobo. Proceedings of the Royal Society B, 292: 20250640.
41. Lewis, L.S., Wessling, E.G., Kano, F., Stevens, J.M.G., Call, J., & Krupenye, C. (2023). Bonobos and chimpanzees remember familiar conspecifics for decades. Proceedings of the National Academy of Sciences, 120(52): e2304903120. doi: 10.1073/pnas.2304903120.
40. Kano, F., Krupenye, C., Hirata, S., Tomonaga, M., & Call, J. (2019). Great apes use self- experience to anticipate an agent’s action in a false belief test. Proceedings of the National Academy of Sciences, 116(42), 20904-20909. doi: 10.1073/pnas.1910095116.
39. Krupenye, C., and Hare, B. (2018). Bonobos prefer individuals that hinder others over those that help. Current Biology. 28(2): 280-286. doi: 10.1016/j.cub.2017.11.061.
38. Krupenye, C.*, Kano, F.*, Hirata, S., Call, J., and Tomasello, M. (2016). Great apes anticipate that other individuals will act according to false beliefs. Science, 354(6308): 110-114. doi: 10.1126/science.aaf8110. -selected by Science as one of the top ten breakthroughs of 2016
Other Publications
37. Morford, J. P, Lewin, P., Mann, R. P., Krupenye, C., & Biro, D. (in press). Collective route memories emerge through differential forgetting of navigational information in homing pigeons. Scientific Reports
36. Bastos, A. P. M.P, Foster, G.G, Wood, P. M., & Krupenye, C. (in press). Do dogs rationally infer the causes of failed actions? PLoS One
35. Lewis, L.S.G, Kano, F., Stevens, J.M.G., DuBois, J.G.G, Call, J., & Krupenye, C. (in press). Do great apes know each other’s names? Probing great ape comprehension of social vocal labels. Animal Behavior and Cognition
34. Townrow, L.G & Krupenye, C. (2025). Reply to Tauzin: How to implement novelty in theory of mind tasks. Proceedings of the National Academy of Sciences, 122 (21) e2503973122.
33. Townrow, L.G & Krupenye, C. (2025). Reply to Royka and Santos: Representing ignorance, and the evolution of theory of mind. Proceedings of the National Academy of Sciences, 122 (20), e2506762122.
32. Krupenye, C., Carvajal, L.G, & Bastos, A. P. M.P (2025). Psychological mechanisms for individual recognition- and anonymous-societies in humans and other animals. Behavioral and Brain Sciences, 48, e65.
31. Bastos, A.P.M., Warren, E., & Krupenye, C. (2024). What evidence can validate a dog training method? Learning & Behavior
30. Krupenye, C. (2024). Divergence in bonobo and chimpanzee social life. Learning & Behavior
29. Westra, E., Fitzpatrick, S., Brosnan, S., Gruber, T., Hobaiter, C., Hopper, L., Kelly, D., Krupenye, C., Luncz, L., Theriault, J., & Andrews, K. (2024). In search of animal normativity: a framework for studying social norms in nonhuman animals. Biological Reviews, 10.1111/brv.13056.
28. Lewis, L.S., & Krupenye, C. (2022). Theory of mind in nonhuman primates. In B.L. Schwartz & M.J. Beran (Eds.) Primate Cognitive Studies. Cambridge University Press.
27. Farrar, B., Altschul, D., Fischer, J., Tennie, C., Ostojic, L., Krupenye, C., & Rodrigo, A. (2022). Replication and reproducibility in primate cognition research. In B.L. Schwartz & M.J. Beran (Eds.) Primate Cognitive Studies. Cambridge University Press.
26. Lewis, L.S., & Krupenye, C. (2022). Eye-tracking as a window into primate social cognition. American Journal of Primatology. doi: 10.1002/ajp.23393.
25. Kano, F., Furuichi, T., Hashimoto, C., Krupenye, C., Leinwand, J.G., Hopper, L.M., Martin, C.F., Otsuka, R., & Tajima, T. (2022). What is so unique about the human eye? Comparative image analysis on the external eye morphology of human and nonhuman great apes. Evolution and Human Behaviour. doi: 10.1016/j.evolhumbehav.2021.12.004
24. Durdevic, K, & Krupenye, C. (2021). Representing knowledge, belief, and everything in between: Representational complexity in humans and other apes. Behavioral and Brain Sciences, 44: e150. doi: 10.1017/S0140525X20001855.
23. Amodio, P., Farrar, B.G., Krupenye, C., Ostojić, L., & Clayton, N. (2021). Little evidence that Eurasian jays protect their caches by responding to cues about a conspecific’s desire and visual perspective. eLife, 10, e69647. doi: 10.7554/eLife.69647.
22. Feldblum, J., Krupenye, C., Bray, J., Pusey, A.E., & Gilby, I.C. (2021). Social bonds provide multiple pathways to reproductive success in wild male chimpanzees. iScience, 24, 102864.
21. Lewis, L.S., Kano, F., Stevens, J.M.G., DuBois, J.G., Call, J., & Krupenye, C. (2021). Bonobos and chimpanzees preferentially attend to familiar members of the dominant sex. Animal Behaviour, 177, 193-206.
20. Hopper, L. M., Gulli, R. A., Howard, L. M., Kano, F., Krupenye, C., Ryan, A., & Paukner, A. (2020). The application of non-invasive, restraint-free eye-tracking methods for use with nonhuman primates. Behavior Research Methods. doi: 10.3758/s13428-020-01465-6.
19. Krupenye, C. (2020). The evolution of mentalizing in humans and other primates. In M. Gilead & K. Ochsner (Eds.) The Neural Basis of Mentalizing: A Social-Cognitive and Affective Neuroscience Perspective. Springer Press.
18. Kano, F., Call, J., & Krupenye, C. (2020). Primates pass dynamically social anticipatory-looking false belief tests. Trends in Cognitive Sciences, doi: 10.1016/j.tics.2020.07.003.
17. Bowie, A., Krupenye, C., Mbonzo, P., Minesi, F., & Hare, B. (2020). Implicit measures help demonstrate the value of conservation education in the Democratic Republic of the Congo. Frontiers in Psychology. doi: 10.3389/fpsyg.2020.00386.
16. Krupenye, C. & Call, J. (2019). Theory of mind in animals: Current and future directions. WIREs Cognitive Science, e1503. read-only link
15. Krupenye, C., Tan, J., & Hare, B. (2018). Bonobos voluntarily hand food to others but not toys or tools. Proceedings of the Royal Society B. 285(1886): 20181536. doi: 10.1098/rspb.2018.1536.
14. Kano, F., Moore, R., Krupenye, C., Hirata, S., Tomonaga, M., & Call, J. (2018). Human ostensive signals do not enhance gaze-following in chimpanzees but do enhance object-oriented attention. Animal Cognition. 21(5): 715-728. doi: https://doi.org/10.1007/s10071-018-1205-z.
13. Krupenye, C., MacLean, E. L., and Hare, B. (2017). Does the bonobo have a (chimpanzee-like) theory of mind? In B. Hare & S. Yamamoto (Eds.) Bonobos: Unique in Mind, Brain, and Behavior. Oxford University Press. DOI: 10.1093/oso/9780198728511.003.0006.
12. Kano, F.*, Krupenye, C.*, Hirata, S., Call, J., and Tomasello, M. (2017). Submentalizing cannot explain belief-based action anticipation in apes. Trends in Cognitive Sciences. 21(9): 633-634. doi: http://dx.doi.org/10.1016/j.tics.2017.06.011.
11. Krupenye, C.*, Kano, F.*, Hirata, S., Call, J., and Tomasello, M. (2017). A test of the submentalizing hypothesis: Apes’ performance in a false belief task inanimate control. Communicative & Integrative Biology. 10(4): e1343771.
10. Krupenye, C. (2017). Guesser-knower paradigm. Encylopedia of Animal Cognition and Behavior. doi: 10.1007/978-3-319-47829-6_1605-1.
9. Kano, F., Krupenye, C., Hirata, S., and Call, J. (2017). Eye tracking uncovered great apes' ability to anticipate that other individuals will act according to false beliefs. Communicative & Integrative Biology. 10(2): e1299836.
8. Reddy, R. B., Krupenye, C., MacLean, E. L., and Hare, B. (2016). No evidence for contagious yawning in lemurs. Animal Cognition, 19(5): 889-898. doi: 10.1007/s10071-016-0986-1.
7. Krupenye, C., Rosati, A. G., and Hare, B. (2016). What’s in a frame? Response to Kanngiesser & Woike (2016). Biology Letters, 12(1): 20150959. doi: 10.1098/rsbl.2015.0959.
6. Foerster, S., McLellan, K., Schroepfer-Walker, K., Murray, C., Krupenye, C., Gilby, I.C., and Pusey, A. (2015). Social bonds in the dispersing sex: partner preferences among adult female chimpanzees. Animal Behaviour, 105: 139-152. doi: 10.1016/j.anbehav.2015.04.012.
5. Krupenye, C., Rosati, A.G., and Hare, B. (2015). Bonobos and chimpanzees exhibit human-like framing effects. Biology Letters, 11(2). doi: 10.1098/rsbl.2014.0527.
4. MacLean, E., Krupenye, C., and Hare, B. (2014). Dogs account for body orientation but not visual barriers when responding to pointing gestures. Journal of Comparative Psychology, 128(3): 285-297. doi: 10.1037/a0035742.
3. Bray, J., Krupenye, C., and Hare, B. (2014). Ring-tailed lemurs (Lemur catta) exploit information about what others can see but not what they can hear. Animal Cognition, 17(3): 735-744. doi: 10.1007/s10071-013-0705-0
2. Rosati, A.G., Herrmann, E., Kaminski, J., Krupenye, C., Melis, A.P., Schroepfer, K., Tan, J., Warneken, F., Wobber, V. & Hare, B. (2012). Assessing the psychological health of captive and wild apes: A response to Ferdowsian et al. (2011). Journal of Comparative Psychology, 127(3): 329-336. doi: 10.1037/a0029144.
1. Gruber, T., Potts, K.B., Krupenye, C., Byrne, M., Mackworth-Young, C., McGrew, W.C., Reynolds, V., & Zuberbuhler, K. (2012). The influence of ecology on chimpanzee cultural behavior: A case study of five Ugandan chimpanzee communities. Journal of Comparative Psychology, 126(4): 446-57. doi: 10.1037/a0028702.