University of Exeter funding: NERC GW4+ DTP PhD studentship

Investment decisions in grey squirrels: the role of future discounting and executive control. PhD in Psychology (NERC GW4+ DTP) Ref: 3707

About the award


Lead Supervisor

Dr. Lisa Leaver, Department of Psychology, College of Life and Environmental Sciences, University of Exeter

Additional Supervisors

Dr. Tim Fawcett, Department of Psychology, College of Life and Environmental Sciences, University of Exeter

Dr. Sean Rands, School of Biological Sciences, University of Bristol

Location: University of Exeter, Streatham Campus, Exeter, EX4 4QJ

This project is one of a number that are in competition for funding from the NERC GW4+ Doctoral Training Partnership (GW4+ DTP).  The GW4+ DTP consists of the GW4 Alliance of research-intensive universities: the University of Bath, University of Bristol, Cardiff University and the University of Exeter plus five unique and prestigious Research Organisation partners: British Antarctic Survey, British Geological Survey, Centre for Ecology & Hydrology, the Natural History Museum and Plymouth Marine Laboratory.  The partnership aims to provide a broad training in the Earth, Environmental and Life sciences, designed to train tomorrow’s leaders in scientific research, business, technology and policy-making. For further details about the programme please see

For eligible successful applicants, the studentships comprises:

  • A stipend for 3.5 years (currently £15,009 p.a. for 2019/20) in line with UK Research and Innovation rates
  • Payment of university tuition fees;
  • A research budget of £11,000 for an international conference, lab, field and research expenses;
  • A training budget of £3,250 for specialist training courses and expenses.
  • Travel and accommodation is covered for all compulsory DTP cohort events
  • No course fees for courses run by the DTP

We are currently advertising projects for a total of 10 studentships at the University of Exeter


Students who are resident in EU countries are eligible for the full award on the same basis as UK residents.  Applicants resident outside of the EU (classed as International for tuition fee purposes) are not eligible for DTP funding. Residency rules are complex and if you have not been resident in the UK or EU for the 3 years prior to the start of the studentship, please apply and we will check eligibility upon shortlisting.

Project Background

Investment behaviour necessarily entails planning for the future. It can take many forms across species, and requires some degree of executive control: investors need to inhibit immediate use of an asset to store it away for a future which is, inevitably, uncertain. Investment decisions provide a window into an animal’s expectations about the future and can be expected to vary with life-history parameters.

Grey squirrels are an ideal model species for studying investment because they are natural and prolific investors, storing nuts during times of plenty to provide a reliable source of food when it is scarce. Squirrels not only decide whether to cache a food item, eat it, or reject it, but caching itself involves a series of predation and pilferage risk trade-offs, which co-vary with food value. As such, investment decisions directly reflect future discounting and ought to be tightly linked to executive control.

Project Aims and Methods

This PhD project will investigate the role of future discounting and executive control in investment decisions made by wild grey squirrels, as part of a long-term field study of squirrels living on the University of Exeter campus. There are two key aims:

1) generate testable predictions of caching behaviour from integrative models that consider both optimal foraging (from the behavioural ecology tradition) and the cognitive mechanisms of discounting and executive control (from experimental/cognitive psychology);

2) develop reliable assays of executive control in squirrels and record their investment decisions and outcomes in terms of successful food recovery, as a real-life measure of the costs and benefits of delaying gratification.

The student will construct mathematical models of adaptive investment behaviour based on executive control and future discounting, to generate predictions that he/she will test by collecting experimental data on wild squirrels in the field. By devising field experiments to verify abstract laboratory tests of executive control, he/she will generate reliable measures of executive control in free-living squirrels and measure the outcomes of their natural investment decisions. There are multiple directions the work could take and we are particularly interested to hear from students who want to play an active role in designing the project.


Marked grey squirrel foraging in the wild



Grey squirrel encountering a test apparatus

Candidate Requirements

The candidate should have experience of working with animals in the lab or field, and an interest in animal cognition, foraging theory and decision-making.


The candidate will be trained in live-trapping and marking of grey squirrels, behavioural observation in the field, video analysis, mathematical modelling and computer programming. They will also be trained in broader subject-relevant skills such as experimental design and statistical analysis.

References / Background reading list

Nevai, A.L., Waite, T.A. & Passino, K. M. (2007) State-dependent choice and ecological rationality. Journal of Theoretical Biology 247; 471–479.

Smulders, T., Gould, K. & Leaver L.A. (2010) Using ecology to guide the study of cognitive and neural mechanisms of different aspects of spatial memory in food-hoarding animals. Philosophical Transactions B. Special edition on food hoarding. 365: 883-900.

Voelter, C. J., Tinklenberg, B., Call, J. & Seed, A.M. (2018) Comparative psychometrics: establishing what differs is central to understanding what evolves. Philosophical Transactions of the Royal Society B. 373, 1756.

Entry requirements

Applicants should have obtained, or be about to obtain, a First or Upper Second Class UK Honours degree, or the equivalent qualifications gained outside the UK.   Applicants with a Lower Second Class degree will be considered if they also have Master’s degree.  Applicants with a minimum of Upper Second Class degree and significant relevant non-academic experience are encouraged to apply.

All applicants would need to meet our English language requirements by the start of the  project

How to apply

In the application process you will be asked to upload several documents.  Please note our preferred format is PDF, each file named with your surname and the name of the document, eg. “Smith – CV.pdf”, “Smith – Cover Letter.pdf”, “Smith – Transcript.pdf”.

  • CV
  • Letter of application outlining your academic interests, prior research experience and reasons for wishing to undertake the project.
  • Transcript(s) giving full details of subjects studied and grades/marks obtained.  This should be an interim transcript if you are still studying.
  • If you are not a national of a majority English-speaking country you will need to submit evidence of your current proficiency in English.

Reference information
You will be asked to name 2 referees as part of the application process, however we will not expect receipt of references until after the shortlisting stage. Your referees should not be from the prospective supervisory team.

If you are shortlisted for interview, please ensure that your two academic referees email their references to the, 7 days prior to the interview dates.  Please note that we will not be contacting referees to request references, you must arrange for them to be submitted to us by the deadline.

References should be submitted by your referees to us directly in the form of a letter. Referees must email their references to us from their institutional email accounts. We cannot accept references from personal/private email accounts, unless it is a scanned document on institutional headed paper and signed by the referee.

All application documents must be submitted in English. Certified translated copies of academic qualifications must also be provided.

The closing date for applications is 1600 hours GMT Monday 6 January 2020.  Interviews will be held between 10 and 21 February 2020.  For more information about the NERC GW4+ DPT please visit

If you have any general enquiries about the application process please email  Project-specific queries should be directed to the lead supervisor.

Data Sharing
During the application process, the University may need to make certain disclosures of your personal data to third parties to be able to administer your application, carry out interviews and select candidates.  These are not limited to, but may include disclosures to:

  • the selection panel and/or management board or equivalent of the relevant programme, which is likely to include staff from one or more other HEIs;
  • administrative staff at one or more other HEIs participating in the relevant programme.

Such disclosures will always be kept to the minimum amount of personal data required for the specific purpose. Your sensitive personal data (relating to disability and race/ethnicity) will not be disclosed without your explicit consent.


Application deadline:6th January 2020
Value:£15,009 per annum for 2019-20
Duration of award:per year
Contact: PGR Enquiries