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Dr Lisa Leaver

Dr Lisa Leaver

Senior Lecturer


 Washington Singer 101


Washington Singer Laboratories, University of Exeter, Perry Road, Prince of Wales Road, Exeter, EX4 4QG, UK


I study the behavioural and cognitive adaptations shaped by selective pressures specific to an animal’s ecology. My research is divided into two main areas; 1) determining how decision-making and behaviour are influenced by risk, and 2) mapping cognitive abilities onto behavioural outcomes in wild animals. I conduct my research in the field using a combination of experimental and observational techniques, focusing primarily on grey squirrels. 


B.A. (Hons) Psychology (Anthropology Minor), McMaster University 1995

Ph. D. Psychology, McMaster University 2000. Thesis entitled “Ecological determinants of foraging and caching behaviour in sympatric heteromyid rodents” Supervised by Prof. Martin Daly, Externally Examined by Prof. David Sherry.

Research group links

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Research interests

My work is focused on exploring behavioural and cognitive adaptations specific to particular aspects of a species’ ecology. I conduct research on the abilities that have been shaped by the selective pressures faced by animals that cache food for use at a later time, mainly squirrels and kangaroo rats. I am interested in questions about the behavioural ecology and cognitive abilities of caching animals and carry out research on the factors affecting their foraging and food caching decisions, risk taking, social learning, categorisation, problem solving, and memory in the wild and in the laboratory.

I am using various techniques to explore the nature of squirrel cognition - for example, I have examined their performance interacting a variety of puzzle boxes in an attempt to explain how they understand and learn about causal relationships.

I am investigating social influences on foraging and food caching decisions made by eastern grey squirrels in the laboratory and in the field. I conduct cognitive and spatial memory experiments in the laboratory, and I look at natural social, foraging and caching behaviour of marked individuals in the field.

I am also investigating individual differences in cost/benefit trade-offs made by animals during foraging and food storing in relation to things such as cues of predation risk, presence of competitors and food value.

Research grants

  • 2019 British Ecological Society (UK)
    Grey squirrels and pine martens: the role of stress in suppressing grey squirrel populations
  • 2007 Great Western Research
    PhD studentship jointly funded by Astra Zenica entitled "Using behaviour to determine welfare and enrichment criteria for aquatic environmental protection research".
  • 2005 EU STREP/NEST
    From Associations to rules in the development of concepts
  • 2001 British Ecological Society (UK)
    The ecological impact of common rats (Rattus norvegicus) on the Isles of Scilly: assessing the value of pest control

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Journal articles

Leaver L, Lea S, Chow P, McLaren I (In Press). Behavioral Flexibility: a review, a model and some exploratory tests. Learning and Behavior
Leaver L, Ford S, Miller C, Yeo M, Fawcett T (In Press). Learning is negatively associated with strength of left/right paw preference in wild grey squirrels (Sciurus carolinensis). Learning and Behavior
Lea SEG, Pothos E, Wills A, Leaver L, Ryan C, Meier C (In Press). Multiple feature use in pigeons’ category discrimination: the influence of stimulus set structure and the salience of stimulus differences. Journal of Experimental Psychology: Animal Learning and Cognition
Leaver L, Wilkinson A (2020). A special issue in honour of Stephen Lea - a true comparative psychologist. Learn Behav, 48(1), 7-8.  Author URL.
Chow PKY, Leaver LA, Wang M, Lea SEG (2017). Touch screen assays of behavioural flexibility and error characteristics in Eastern grey squirrels (Sciurus carolinensis). Animal Cognition, 20(3), 459-471.
Leaver LA, Jayne K, Lea SEG (2016). Behavioral flexibility versus rules of thumb: how do grey squirrels deal with conflicting risks?. Behavioral Ecology, 28(1), 186-192.
Chow PKY, Lea SEG, Leaver LA (2016). How practice makes perfect: the role of persistence, flexibility and learning in problem solving efficiency. Animal Behaviour, 273-283. Abstract.
Jayne K, Lea SEG, Leaver LA (2015). Behavioural responses of Eastern grey squirrels, <i>Sciurus carolinensis</i>, to cues of risk while foraging. BEHAVIOURAL PROCESSES, 116, 53-61.  Author URL.
Chow PKY, Leaver LA, Wang M, Lea SEG (2015). Serial Reversal Learning in Gray Squirrels: Learning Efficiency as a Function of Learning and Change of Tactics. JOURNAL OF EXPERIMENTAL PSYCHOLOGY-ANIMAL LEARNING AND COGNITION, 41(4), 343-353.  Author URL.
Farmer HL, Plowman AB, Leaver LA (2011). Role of vocalisations and social housing in breeding in captive howler monkeys (Alouatta caraya). Applied Animal Behaviour Science, 134(3-4), 177-183. Abstract.
Hopewell LJ, Leaver LA, Lea SEG, Wills AJ (2010). Grey squirrels (Sciurus carolinensis) show a feature-negative effect specific to social learning. Anim Cogn, 13(2), 219-227. Abstract.  Author URL.
Smulders T, Gould K, Leaver LA (2010). Using ecology to guide the study of cognitive and neural mechanisms of different aspects of spatial memory in food-hoarding animals. Philosophical Transactions of the Royal Society B, 365, 883-900. Abstract.
Wills AJ, Lea SEG, Leaver LA, Osthaus B, Ryan CME, Suret MB, Bryant CML, Chapman SJA, Millar L (2009). A comparative analysis of the categorization of multidimensional stimuli: I. unidimensional classification does not necessarily imply analytic processing; evidence from pigeons (Columba livia), squirrels (Scurius carolinensis) and humans (Homo sapiens). Journal of Comparative Psychology, 123(4), 391-405. Abstract.
Lea SEG, Wills AJ, Leaver LA, Ryan CME, Bryant CML, Millar L (2009). A comparative analysis of the categorization of multidimensional stimuli: II. Strategic information search in humans (Homo sapiens) but not in pigeons (Columba livia). Journal of Comparative Psychology, 123(4), 406-420. Abstract.
Gilbert-Norton LB, Leaver LA, Shivik JA (2009). The effect of randomly altering the time and location of feeding on the behaviour of captive coyotes (Canis latrans). APPL ANIM BEHAV SCI, 120(3-4), 179-185. Abstract.
Jule KR, Lea SEG, Leaver LA (2009). Using a behaviour discovery curve to predict optimal observation time: captive red pandas (Aulurus fulgens) as a case study. Behaviour, 146, 1531-1542. Abstract.
Hopewell LJ, Leaver LA, Lea SEG (2008). Effects of Competition and Food Availability on Travel Time in Scatter-hoarding Grey Squirrels (Sciurus carolinensis). Behavioral Ecology(19), 1143-1149.
Hopewell LJ, Leaver LA (2008). Evidence of social influences on cache-making by grey squirrels (Sciurus carolinensis). Ethology, 114, 1061-1068.
Jule K, Leaver L, Lea SEG (2008). The effects of captive experience on reintroduction survival in carnivores: a review and analysis. Biological Conservation, 141, 355-363. Abstract.
Leaver LA, Caldwell C, Hopewell L, Mallarky L (2007). Audience effects of food caching in grey squirrels (Sciurus carolinensis): evidence for pilferage avoidance strategies. Animal Cognition, 10(1), 23-27.
Hopewell LJ, Rossiter, R. Blower, E. Leaver, L. Goto K (2005). Grazing and vigilance by Soay Sheep on Lundy Island: influence of group size, terrain and the distribution of vegetation. Behavioural Processes, 70, 186-193.  Author URL.
Leaver LA (2004). Effects of food value, predation risk and pilferage on the caching decisions of Dipodomys merriami. Behavioral Ecology, 15(5), 729-734.
Leaver LA, Daly M (2003). Effects of predation risk on selectivity in heteromyid rodents. Behavioural Processes, 64(1), 71-75.
Leaver LA, Daly M (2001). Food caching and differential cache pilferage: a field study of co-existence of sympatric kangaroo rats and pocket mice. Oecologia, 128(4), 577-584.
Leaver LA, Daly, M. (1998). Effects of food preference on scatter hoarding by kangaroo rats (Dipodomys merriami). Behaviour, 135, 823-832.  Author URL.


Leaver LA, Jayne K (2015). Strategic decisions made by small mammals during scatter hoarding, cache recovery and cache pilferage. In Shuttleworth C, Lurz P, Hayward M (Eds.) Red Squirrels: Ecology, Conservation & Management in Europe, European Squirrel Initiative, 51-65.


Wilkes L, Leaver L, Sloman K, Readman G, Williams T, Owen S, Wilson R (2009). Development of enrichment criteria for zebrafish(<i>Danio rerio</i>) used in laboratory studies.  Author URL.
Landin J, Wilson R, Owen S, Williams T, Readman G, Sloman K, Leaver L (2009). Welfare and enrichment criteria for fish used in research.  Author URL.

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External Engagement and Impact

Administrative responsibilities

2011 – 2013 - External Examiner for the MRes in Animal Behaviour at Newcastle University

2011 – External Assessor for the BSc in Animal Behaviour at Cornwall College, Newquay.

Media Coverage

My squirrel caching research was featured on “Autumnwatch”, BBC2.  Aired Tuesday October 30, 2012.

Interviewed on BBC Radio Devon, discussing squirrel intelligence. Tuesday October 30, 2012.

Guest on Countywide Breakfast with Matt Woodley. Aired Wednesday July 29, 2009. BBC Radio Devon

“May (or may not) contain nuts”. Planet Earth online, July 28, 2009.

“They know they’re being watched”. New Scientist. January 19, 2008.

David Attenborough’s ‘The Life of Mammals’ Series episode entitled ‘The Seed Predators’. Aired Wednesday December 11, 2002. BBC1.

Bill Oddie Goes Wild On the Isles of Scilly. Aired Friday January 4, 2002. BBC2.

Workshops organised

Member of the organising committe for the 16th congress of the International Society for Behavioral Ecology, University of Exeter, July 28- August 3, 2016.

Co-organiser of Integrative Biology of Scatter Hoarding: Ecology, Psychology and Neuroscience, Newcastle University, 9 August 2013.

Member of the organising committee for Association for the Study of Animal Behaviour Easter Meeting, University of Exeter, April 7-9, 2010.

Member of the organising committee for Integrative Biology of Scatter Hoarding: Ecology, Psychology and Neuroscience, Cornell University (Ithaca, NY), August 8-9, 2008.

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My teaching focusses on animal behaviour, the evolutionary basis of behaviour, animal cognition and human evolutionary psychology and I lead the MSc in Animal Behaviour annual field trip to Lundy Island.



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Supervision / Group

Postgraduate researchers

  • Yavanna Burnham
  • Ingerid Helgestad
  • Robert Kelly


  • Pizza Ka Yee Chow
  • Rebecca Cummins
  • Holly Farmer
  • Lucy Hopewell
  • Kimberley Jayne
  • Kristen Jule
  • Jenny Landin
  • Helen Matthews
  • Louise Millar
  • Kirsten Pullen

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