Staff profile

I am a comparative and developmental psychologist with expertise in primatology. I study and compare great apes and young children in order to investigate the evolutionary and developmental basis of hominid social cognition and behavior. My main interests are the development & evolution of social cognition and communication, focussing on empathy, language and social learning. 

 Research Interests 

Evolutionary & Developmental Perspectives on:

• Social cognition and communication
• Empathy & socio-emotional processes
• Language evolution
• Child gesture
• Social learning and cultural cognition
• Primate vocal communication (including referential communication & call combinations) 

I am a specialist in great ape behaviour, with particular expertise in bonobos, our closest living relatives. I have experience studying bonobos in the wild, captivity and semi-captivity, including Lola ya Bonobo Sanctuary - where I also act as a long-term Scientific Consultant. 

I conducted my doctoral work with Prof Klaus Zuberbuhler (University of St Andrews) investigating bonobo vocal communication and language evolution. I conducted post-doctoral work with Prof Frans de Waal (Emory University, USA) examining empathy development in sanctuary-living bonobos. I conducted post-doctoral research in the Dept of Comparative Cognition, University of Neuchatel (Switzerland) studying vocal communication in wild bonobos at Lui Kotale, DR Congo (MPI-EVA).

I completed a Marie Curie Research Fellowship at the University of Birmingham with Dr Claudio Tennie examining the evolution & development of imitation and language with great apes and children.

Available PhD and MRes research projects

I would welcome applications from people interested in Comparative and Developmental Psychology, more specifically on the origins of communication, empathy and social cognition. I would be happy to supervise both experimental and observational projects working with human children (UK and cross-culturally) and primates. I have a good network of contacts to arrange access to a variety of primates, including wild, sanctuary-living and captive bonobos. For those wishing to embark on a fieldwork PhD study, experience of travel and work in third world countries is a real advantage.

Please contact me if you are interested in making an application

• Primate Conservation
• Comparative Affective Science
• Social cognition
• Evolution of language and culture
• Evolution and development of empathy
• Primate behaviour
• Science outreach - primatology
• Development of communication and social learning
• Primate vocal communication
• Animal cognition
• Child gesture development
• Applications to primate conservation

• Developmental Science

• BBC Article I contributed to on Science of Smiles
• Comments for media article http://www.wunc.org/post/whats-mine-yours-sort-bonobos-and-tricky-evolutionary-roots-sharing#stream/0
• Comments for press coverage https://www.bbc.co.uk/news/science-environment-45114322
• Coverage in The Guardian about research on bonobo calls and speech
• Coverage of imitation paper in Child Development
• Coverage of imitation paper in Child Development
• Discussing comparative cognition in IB Times
• Invited Guest on Radio 4 'Inside Science' to discuss bonobo calls
• Media article from New Scientist

Available for media contact about:

• Primate behaviour: I am a comparative and devlopmental psychologist interested in origins of human behaviour from developmental and evolutionary perspectives. I study great ape communication and behaviour in order to ask questions about human evolution.
• Conflict and resolution: I am interested in the evolution of conflict resolution by studying conflict and post-conflict resolution in great apes. This extends to studying consolation, a marker of empathy.
• Aggression: I am interested in the extent to which aggression and peaceful coexistence shape the societies of our closest living relatives, chimpanzees and bonobos
• Wildlife: I am passionate about supporting the conservation of primates, and am actively assist bonobo conservation in conjucntion with Lola ya Bonobo Sanctuary, the worlds only bonobo sanctuary
• Linguistics & Language: My research interests include the evolution of language and communication using insights from non-human primates
• Human biology and development: Development of communication, with a focus on child gesture, and socio-emotional competence in young chidren
• Psychology: Development of communication, with a focus on child gesture, and socio-emotional competence in young chidren
• Evolution:
• Primate behaviour:
• General issues:
• Human impact:
• Wildlife:
• Africa:
• Human biology and development:
• Psychology:
• Aggression:
• Relationships:
• Stress:
• Science & Technology:
• People: Evolution and Biology:
• The Earth: Animals:

Chapter in book

• Clay, Z., Webb, C. E., Romero, T. & de Waal F. B. M. (2022). Comparative Perspectives of Empathy Development: Insights from chimpanzees and bonobos. In The Oxford Handbook of Emotional Development. Dukes, D., Walle, E. & Samson, A. Oxford: Oxford University Press. 277-290.
• Clay, Zanna, Palagi, Elisabetta & de Waal, Frans B.M. (2018). Ethological Approaches to Empathy in Primates. In Neuronal correlates of empathy – from rodent to man. Meyza, Ksenia & Knapska, Ewelina Elsevier.
• Clay, Zanna Palagi, Elisabetta & de Waal, Frans B.M. (2018). Ethological Approaches to Empathy in Primates. In Neuronal Correlates of Empathy: From Rodent to Man. Ksenia Meyza & Ewelina Knapska Academic Press. 53-66.
• Clay, Zanna & Genty, Emilie (2017). Natural communication in bonobos: Insights into social awareness and the evolution of language. In Bonobos: Unique in Mind Brain and Behavior. Hare, Brian & Yamamoto, Shinya Oxford: Oxford University Press.
• Clay, Zanna & Zuberbuhler, Klaus (2014). Vocal communication and social awareness in chimpanzees and bonobos. In The Social Origins of Language. Dor, Daniel, Knight, Chris & Lewis, Jerome Oxford University Press.
• Clay, Zanna & Iacoboni, Marco (2011). Mirroring Fictional Others. In The Aesthetic Mind, Philosophy and Psychology. Schellekens, Elisabeth & Goldie, Peter Oxford University Press.

Journal Article

• Schamberg, Isaac, Clay, Zanna, Townsend, Simon W. & Surbeck, Martin (2023). Between-group variation in production of pant-grunt vocalizations by wild bonobos (Pan paniscus). Behavioral Ecology and Sociobiology 77(1): 14, 1-12.
• Kim, Y., Vlaeyen, J., Heesen, R., Clay, Z. & Kret, M. (2022). The Association Between the Bared-Teeth Display and Social Dominance in Captive Chimpanzees (Pan troglodytes). Affective Science 3(4): 749-760.
• Heesen, R., Austry, D.A., Upton, Z. & Clay, Z. (2022). Flexible signalling strategies by victims mediate post-conflict interactions in bonobos. Philosophical Transactions of the Royal Society B: Biological Sciences 377(1860).
• Vlaeyen, Jolinde M. R., Heesen, Raphaela, Kret, Mariska E., Clay, Zanna, Bionda, Thomas & Kim, Yena (2022). Bared-teeth displays in bonobos (Pan paniscus): An assessment of the power asymmetry hypothesis. American Journal of Primatology 84(9): e23419.
• Demuru, E, Clay, Z & Norscia, I (2022). What makes us apes? The emotional building blocks of intersubjectivity in hominids. Ethology Ecology & Evolution 34(3): 220-234.
• Derry, T, Clay, Z, Dahl, CD, Zuberbühler, J, Davila-Ross, M & Dezecache, G (2022). Vocal functional flexibility: what it is and why it matters. Animal Behaviour 186: 93-100.
• Kavanagh, Eithne, Street, Sally E., Angwela, Felix O., Bergman, Thore J., Blaszczyk, Maryjka B., Bolt, Laura M., Briseño-Jaramillo, Margarita, Brown, Michelle, Chen-Kraus, Chloe, Clay, Zanna, Coye, Camille, Thompson, Melissa Emery, Estrada, Alejandro, Fichtel, Claudia, Fruth, Barbara, Gamba, Marco, Giacoma, Cristina, Graham, Kirsty E., Green, Samantha, Grueter, Cyril C., Gupta, Shreejata, Gustison, Morgan L., Hagberg, Lindsey, Hedwig, Daniela, Jack, Katharine M., Kappeler, Peter M., King-Bailey, Gillian, Kuběnová, Barbora, Lemasson, Alban, Inglis, David MacGregor, Machanda, Zarin, MacIntosh, Andrew, Majolo, Bonaventura, Marshall, Sophie, Mercier, Stephanie, Micheletta, Jérôme, Muller, Martin, Notman, Hugh, Ouattara, Karim, Ostner, Julia, Pavelka, Mary S. M., Peckre, Louise R., Petersdorf, Megan, Quintero, Fredy, Ramos-Fernández, Gabriel, Robbins, Martha M., Salmi, Roberta, Schamberg, Isaac, Schoof, Valérie A. M., Schülke, Oliver, Semple, Stuart, Silk, Joan B., Sosa-Lopéz, J. Roberto, Torti, Valeria, Valente, Daria, Ventura, Raffaella, van de Waal, Erica, Weyher, Anna H., Wilke, Claudia, Wrangham, Richard, Young, Christopher, Zanoli, Anna, Zuberbühler, Klaus, Lameira, Adriano R. & Slocombe, Katie (2021). Dominance style is a key predictor of vocal use and evolution across nonhuman primates. Royal Society Open Science 8(7): 210873.
• Williams, Lisa A., Brosnan, Sarah F. & Clay, Zanna (2020). Anthropomorphism in comparative affective science: Advocating a mindful approach. Neuroscience & Biobehavioral Reviews 115: 299-307.
• Kret,M.E,, Prochazkova, E,, Sterc, E. H. M, & Clay, Z. (2020). Emotional expressions in human and non-human great apes. Neuroscience & Biobehavioral Reviews 115: 378-395.
• Grawunder, S., Crockford, C., Clay, Z., Kalan, A. K., Stevens, J. M. G., Stoessel, A. & Hohmann, G. (2018). Higher fundamental frequency in bonobos is explained by larynx morphology. Current Biology 28(20): R1188-R1189.
• Clay, Z., Over, H. & Tennie, C. (2018). What drives young children to over-imitate? Investigating the effects of age, context, action type, and transitivity. Journal of Experimental Child Psychology 166: 520-534.
• Clay, Z. & Tennie, C. (2018). Is Overimitation a Uniquely Human Phenomenon? Insights From Human Children as Compared to Bonobos. Child Development 89(5): 1535-1544.
• Schamberg, Isaac, Cheney, Dorothy L., Clay, Zanna, Hohmann, Gottfried & Seyfarth, Robert M. (2017). Bonobos use call combinations to facilitate inter-party travel recruitment. Behavioral Ecology and Sociobiology 71(4): 75.
• Schamberg, Isaac, Cheney, Dorothy L., Clay, Zanna, Hohmann, Gottfried & Seyfarth, Robert M. (2016). Call combinations, vocal exchanges and interparty movement in wild bonobos. Animal Behaviour 122: 109-116.
• Gruber, Thibaud & Clay, Zanna (2016). A Comparison Between Bonobos and Chimpanzees: A Review and Update. Evolutionary Anthropology: Issues, News, and Reviews 25(5): 239-252.
• Clay, Zanna, Ravaux, Lucie, de Waal, Frans B. M. & Zuberbühler, Klaus (2016). Bonobos (Pan paniscus) vocally protest against violations of social expectations. Journal of Comparative Psychology 130(1): 44-54.
• Clay, Zanna, de Waal, Frans B.M. & Furuichi, Takeshi (2016). Obstacles and catalysts to peaceful coexistence in chimpanzees and bonobos. Behaviour 153(9-11): 1293-1330.
• Clay, Zanna, Archbold, Jahmaira & Zuberbühler, Klaus (2015). Functional flexibility in wild bonobo vocal behaviour. PeerJ 3: e1124.
• Scarantino, Andrea & Clay, Zanna (2015). Contextually variable signals can be functionally referential. Animal Behaviour 100: e1-e8.
• Clay, Zanna & de Waal, Frans B.M. (2015). Sex and strife: post-conflict sexual contacts in bonobos. Behaviour 152(3-4): 313-334.
• Clay, Zanna, Pople, Sally, Hood, Bruce & Kita, Sotaro (2014). Young Children Make Their Gestural Communication Systems More Language-Like: Segmentation and Linearization of Semantic Elements in Motion Events. Psychological Science 25(8): 1518-1525.
• Genty, Emilie, Clay, Zanna, Hobaiter, Catherine & Zuberbühler, Klaus (2014). Multi-Modal Use of a Socially Directed Call in Bonobos. PLoS ONE 9(1): e84738.
• Clay, Z. & de Waal, F. B. M. (2013). Development of socio-emotional competence in bonobos. Proceedings of the National Academy of Sciences 110(45): 18121-18126.
• Clay, Zanna & de Waal, Frans B. M. (2013). Bonobos Respond to Distress in Others: Consolation across the Age Spectrum. PLoS ONE 8(1): e55206.
• Clay, Zanna & Zuberbühler, Klaus (2012). Communication during sex among female bonobos: effects of dominance, solicitation and audience. Scientific Reports 2: 291.
• Clay, Zanna, Smith, Carolynn L. & Blumstein, Daniel T. (2012). Food-associated vocalizations in mammals and birds: what do these calls really mean? Animal Behaviour 83(2): 323-330.
• Clay, Zanna & Zuberbühler, Klaus (2011). The Structure of Bonobo Copulation Calls During Reproductive and Non-Reproductive Sex. Ethology 117(12): 1158-1169.
• Clay, Zanna & Zuberbühler, Klaus (2011). Bonobos Extract Meaning from Call Sequences. PLoS ONE 6(4): e18786.
• Clay, Z., Pika, S., Gruber, T. & Zuberbuhler, K. (2011). Female bonobos use copulation calls as social signals. Biology Letters 7(4): 513.
• Gruber, Thibaud, Clay, Zanna & Zuberbühler, Klaus (2010). A comparison of bonobo and chimpanzee tool use: evidence for a female bias in the Pan lineage. Animal Behaviour 80(6): 1023-1033.
• Clay, Zanna & Zuberbühler, Klaus (2009). Food-associated calling sequences in bonobos. Animal Behaviour 77(6): 1387-1396.