Clark, H., & Leavens, D. A. (2021) The performance of domestic dogs (Canis lupus familiaris) on two versions of the Object Choice Task. Animal Cognition. (Published online in advance of print.)

Leavens, D. A. (2021) The referential problem space revisited: An ecological hypothesis of the evolutionary and developmental origins of pointing. Wiley Interdisciplinary Reviews: Cognitive Science. DOI:10.1002/wcs.1554

Leavens, D. A., & Bard, K. A. (In press). Primate cognition in captivity. In A. Lock, C. Sinha, and N. Gontier (Eds.), Oxford handbook of human symbolic evolution.  Oxford: Oxford University Press.


Bates LA (2020) Cognitive abilities in elephants. In: The Cambridge Handbook of Evolutionary Perspectives on Human Behaviour. Eds. L. Workman, W. Reader & J. Barkow. Cambridge University Press, Cambridge. Pp 14-22.

Clark, H., Flack, Z. M., & Leavens, D. A. (2020) Changes in the referential problem space of infants and toddlers (Homo sapiens): Implications for cross-species comparisons. Journal of Comparative Psychology 134: 330-340.

Humphreys, T., Stringer, F., Proops, L. & McComb, K. (2020) Slow blink eye closure in shelter cats Is related to quicker adoption. Animals 2020, 10, 2256.

Humphrey, T., Proops, L., Forman, J., Spooner, R. & McComb, K. (2020) The role of cat eye narrowing movements in cat-human communication. Scientific Reports 10:16503.

van de Water A, Henley M, Bates LA & Slotow R (2020) Future of Thailand’s captive elephants: Commentary on Baker & Winkler on Elephant Rewilding. Animal Sentience 28 (18) 326.   


Bard, K. A., & Leavens, D. A. (2019, June) A menagerie of human concepts: A review of M. Tomasello, Becoming human: A theory of ontogeny. The Psychologist 32: 65.

Bates LA & Byrne RW (2019) The Evolution of Intelligence: Reconstructing the Pathway to the Human Mind. In: The Cambridge Handbook of Intelligence, 2nd Edition. Ed. R. Sternberg. Cambridge University Press, Cambridge. Pp 428-450.

Clark, H., & Leavens, D. A. (2019). Testing dogs in ape-like conditions: The effect of a barrier on dogs’ performance on the object choice task. Animal Cognition 22: 1063-1072.

Clark, H., Elsherif, M., & Leavens, D. A. (2019) Ontogeny vs. phylogeny in Primate/Canid comparisons: A meta-analysis of the object choice task. Neuroscience and Biobehavioral Reviews 105: 178-189.

Leavens, D. A. (2019, October 1) The pointing ape. Aeon.

Leavens, D. A., Bard, K. A., & Hopkins, W. D. (2019) The mismeasure of ape social cognition. Animal Cognition 22: 487-504.


Bates LA (2018) Elephants – Studying Cognition in the African Savannah. In: Field and Laboratory Methods in Animal Cognition. Eds. N. Bueno-Guerra & F. Amici. Cambridge University Press, Cambridge.

Flack, Z., & Leavens, D. A. (2018) Communication and developmental milestones. In T. K. Shackelford & V. E. Weekes-Shackelford (Eds.), Encyclopedia of evolutionary psychological science. New York: Springer.

Flack, Z., Naylor, M., & Leavens, D. A. (2018) Pointing to visible and invisible targets. Journal of Nonverbal Behavior 42: 221-236.

Garai ME, Bates LA, Bertschinger H, Delsink A, Pretorius Y, Selier J, Zitzer HR (2018) Non-lethal elephant population control methods: Summary of the first workshop of the Elephant Specialist Advisory Group of South Africa. Bothalia – African Biodiversity and Conservation 48 (2) a2357.   

Krause, M. A., Udell, M. A. R., Leavens, D. A., & Skopos, L. (2018) Animal pointing: Changing trends and findings from 30 years of research. Journal of Comparative Psychology 132: 326-345.

Leavens, D. A. (2018) The cognitive implications of intentional communication: A multi-faceted mirror. In L. Di Paolo, F. Di Vincenzo, & F. De Petrillo (Eds.), Evolution of primate social cognition (pp. 59-77). New York: Springer.

Leavens, D. A. (2018) William Hopkins. In J. Vonk & T. K. Shackelford (Eds.), Encyclopedia of animal cognition and behavior. New York: Springer.

Pretorius Y, Garai M & Bates LA (2018) The status of African elephant Loxodonta africana populations in South Africa. Oryx. doi:10.1017/S0030605317001454.

Proops L., Grounds K., Smith A.V. & McComb K. (2018) Animals remember previous facial expressions that specific humans have exhibited. Current Biology 28, 1428–1432.e4

Smith A.V., Proops L., Grounds K., Wathan J., Scott S. & McComb K. (2018) Domestic horses (Equus caballus) differentiate negative from positive emotion in human nonverbal vocalisations. Scientific Reports 8:13052 (2018).


Benson-Amram, S., Gilfillan, G. & McComb, K. (2017)  Numerical assessment in the wild: insights from social carnivores  Philosophical Transactions R. Soc.: Biological Sciences, 373: 20160508.

Garai ME, Bates LAPretorius Y, Hofmeyr M, Henley M, Selier J (2017) Understanding Elephants: Guidelines for safe and enjoyable elephant viewing. Struik Nature: South Africa.

Gilfillian, Geoffrey D., Vitale, Jessica D. T., McNutt, J. Weldon, McComb, Karen (2017) Spontaneous discrimination of urine odours in wild African lions (Panthera leoAnimal Behaviour, 126 pp. 177-185. ISSN 0003-3472.

Leavens, D. A., & Clark, H. (2017). Joint attention. In J. Vonk & T. K. Shackelford (Eds.), Encyclopedia of animal cognition and behavior. New York: Springer. DOI: 10.1007/978-3-319-47829-6_1075-1

Smith A.V., Wilson C., McComb K. & Proops L. (2017) Domestic horses (Equus caballus) prefer to approach humans displaying a submissive rather than a dominant body posture. Animal Cognition 21, 307-312.


Gilfillan, G, Vitale, J., McNutt, J.W. & McComb, K (2016) Cross-modal individual recognition in wild African Lions. Biology Letters 12: 20160323.

Leavens, D. A., & Bard, K. A. (2016) Quick guide: Tickling. Current Biology 26: R91-R93.

Leavens, D. A., & Hopkins, W. D. (2016) Primate gestures. In T. K. Shackelford & V. E. Weekes-Shackelford (Eds.), Encyclopedia of evolutionary psychological science. New York: Springer.

Smith, A., Proops, L., Grounds, K., Wathan, J. & McComb, K. (2016) Functionally relevant responses to human facial expressions of emotion in the domestic horse (Equus caballus). Biology Letters 12: 20150907.

Wathan, J., Proops, L., Ground, K. & McComb, K. (2016) Horses discriminate between facial expressions of conspecifics. Scientific Reports 6:38322.


Leavens, D. A., Reamer, L. A., Mareno, M. C., Russell, J. L., Wilson, D., Schapiro, S. J., & Hopkins, W. D. (2015) Distal communication by chimpanzees (Pan troglodytes): Evidence for common ground? Child Development 86: 1623-1638.

Wathan, J., Burrows, A, Waller, B. & McComb, K. (2015) EquiFACS: The Equine Facial Action Coding System. PLoS ONE 10(8): e0131738.


Bard, K. A., & Leavens, D. A. (2014) The importance of development for comparative primatology. Annual Review of Anthropology 43: 183-200.

Bard, K. A., Bakeman, R., Boysen, S. T., & Leavens, D. A. (2014) Emotional engagements predict and enhance social cognition in young chimpanzees.  Developmental Science 17: 682-696.

Froese, T., & Leavens, D. A. (2014) The direct perception hypothesis: Perceiving the intention of another’s action hinders its precise imitation. Frontiers in Psychology 5(65): 1-15.

Hobaiter, C., Leavens, D. A., & Byrne, R. W. (2014). Deictic gesturing in wild chimpanzees? Some possible cases. Journal of Comparative Psychology 128:, 82-87. [Reprinted in French as Hobaiter, C., Leavens, D. A., & Byrne, R. W. (2016). Gestes déictiques chez les chimpanzés sauvages (Pan troglodytes)? Quelques cas possibles. Enfance 2016(4): 405-417.]

Leavens, D. A. (2014) The plight of the sense-making ape. In M. Cappuccio & T. Froese (Eds.), Enactive cognition at the edge of sense-making (pp. 81-104). Basingstoke, U.K.: Palgrave Macmillan.

Leavens, D. A., Sansone, J., Burfield, A. R., Lightfoot, S., O'Hara, S., & Todd, B. K. (2014). Putting the 'joy' in joint attention: Affective-gestural synchrony by parents who point for their babies. Frontiers in Psychology 5(879): 1-7. [Reprinted in E. Di Paolo & H. De Jaegher (eds.). (2015) Towards an embodied science of intersubjectivity: Widening the scope of social understanding research (pp. 171-177). Lausanne: Frontiers Media. doi: 10.3389/978-2-88919-529-9.]

Leavens, D. A., Taglialatela, J. P., & Hopkins, W. D. (2014) From grasping to grooming to gossip: Innovative use of chimpanzee signals in novel environments supports both vocal and gestural theories of language origins. In M. Pina & N. Gontier (Eds.), The evolution of social communication in primates: A multidisciplinary approach (pp. 179-194). New York: Springer.

Lyn, H., Russell, J. L., Leavens, D. A., Bard, K. A., Boysen, S. T., Schaeffer, J., & Hopkins, W. D. (2014) Apes communicate about absent and displaced objects: Methodology matters. Animal Cognition 17: 85-94.

McComb, K. Shannon, G., Sayialel, K. & Moss, C. (2014) Elephants can determine ethnicity, gender, and age from acoustic cues in human voices PNAS 111(14), 5433–5438.

Meguerditchian, A., Taglialatela, J. P., Leavens, D. A., & Hopkins, W. D. (2014) Why vocal production of atypical sounds in apes and its cerebral correlates have a lot to say about the origin of language. Behavioral and Brain Sciences 37: 565-566.

Ratcliffe, V.F., McComb, K. & Reby, D. (2014) Cross-modal discrimination of human gender by domestic dogs. Animal Behaviour 91, 127–135.

Wathan, J. & McComb, K. (2014) The eyes and ears are visual indicators of attention in domestic horses.  Current Biology 24, R1-R2.