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Durham University

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Staff Profile

Dr Zanna Clay, PhD

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Assistant Professor in the Department of Psychology
Telephone: +44 (0) 191 33 49114
Room number: RH009

Contact Dr Zanna Clay (email at


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

Research Groups

Department of Psychology

Research Interests

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

Teaching Areas

  • Advanced Developmental Psychology Review
  • Classic Papers
  • Current Issues in Developmental Psychology & Psychopathology (MSc)
  • Introduction to Psychology 2
  • Research Methods in Psychology

Selected Publications

Chapter in book

  • 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, Palagi, Elisabetta & de Waal, Frans B.M. (2017). Ethological Approaches to Empathy in Primates. In Neuronal correlates of empathy – from rodent to man. Meyza, Ksenia & Knapska, Ewelina Elsevier.
  • 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

  • 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. & Tennie, C. (2018). Is Overimitation a Uniquely Human Phenomenon? Insights From Human Children as Compared to Bonobos. Child Development 89(5): 1535-1544.
  • 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.
  • 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.
  • 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.
  • 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.
  • 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.
  • Scarantino, Andrea & Clay, Zanna (2015). Contextually variable signals can be functionally referential. Animal Behaviour 100: e1-e8.
  • Clay, Zanna, Archbold, Jahmaira & Zuberbühler, Klaus (2015). Functional flexibility in wild bonobo vocal behaviour. PeerJ 3: e1124.
  • Clay, Zanna & de Waal, Frans B.M. (2015). Sex and strife: post-conflict sexual contacts in bonobos. Behaviour 152(3-4): 313-334.
  • 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, 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.
  • 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, 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 & 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). 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.
  • Clay, Zanna & Zuberbühler, Klaus (2011). The Structure of Bonobo Copulation Calls During Reproductive and Non-Reproductive Sex. Ethology 117(12): 1158-1169.
  • 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.

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Media Contacts

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: