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Award details

University of Exeter funding: NERC GW4+ DTP PhD studentship

Investment decisions in grey squirrels: The role of future discounting and executive control. Psychology PhD studentship (NERC GW4+ DTP funded) Ref: 4027

About the award


Lead Supervisor

Dr Lisa Leaver, University of Exeter.

Additional Supervisors

Dr Tim Fawcett, University of Exeter.
Dr Sean Rands, University of Bristol.

Location: Streatham Campus, University of Exeter, Exeter, Devon.

This project is one of a number that are in competition for funding from the NERC Great Western Four+ Doctoral Training Partnership (GW4+ DTP).  The GW4+ DTP consists of the Great Western Four alliance of the University of Bath, University of Bristol, Cardiff University and the University of Exeter plus five Research Organisation partners:  British Antarctic Survey, British Geological Survey, Centre for Ecology and Hydrology,  the Natural History Museum and Plymouth Marine Laboratory.  The partnership aims to provide a broad training in earth and environmental sciences, designed to train tomorrow’s leaders in earth and environmental science. For further details about the programme please see

For eligible successful applicants, the studentships comprises:

  • An stipend for 3.5 years (currently £15,285 p.a. for 2020-21) 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.
  • Up to £750 for travel and accomodation for compulsory cohort events.

Project details


Marked grey squirrel foraging in the wild



Grey squirrel encountering a test apparatus


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 or eat a food item, 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) 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.

2) 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);

By devising field experiments to verify abstract laboratory tests of executive control, the student will generate reliable measures of executive control in free-living squirrels and measure the outcomes of their natural investment decisions. 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. 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.

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.

Background reading and references:

McNamara, J.M., Trimmer, P.C. & Houston, A.I. (2012) The ecological rationality of state-dependent valuation. Psychological Review. 119, 114–119.

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.

Useful Links:

For information relating to the research project please contact the lead Supervisor via email:;

Prospective applicants:

For information about the application process please contact the Admissions team via

Each research studentship project advertisement has an ‘Apply Now’ button linking to an application portal. Please note that applications received via other routes including a standard programme application route will not be considered for the studentship funding.

How to Apply:

The application deadline is Friday 8 January 2021 at 2359 GMT. Interviews will take place from 8th to 19th February 2021. For more information about the NERC GW4+ Doctoral Training Partnership please visit


NERC GW4+ DTP studentships are open to UK and Irish nationals who, if successful in their applications, will receive a full studentship including payment of university tuition fees at the home fees rate.

A limited number of full studentships are also available to international students which are defined as EU (excluding Irish nationals), EEA, Swiss and all other non-UK nationals.  For further details please see the NERC GW4+ website.

Those not meeting the nationality and residency requirements to be treated as a ‘home’ student may apply for a limited number of full studentships for international students. Although international students are usually charged a higher tuition fee rate than ‘home’ students, those international students offered a NERC GW4+ Doctoral Training Partnership full studentship starting in 2021 will only be charged the ‘home’ tuition fee rate (which will be covered by the studentship). International applicants need to be aware that you will have to cover the cost of your student visa, healthcare surcharge and other costs of moving to the UK to do a PhD. More information on this is available from the universities you are applying to (contact details are provided in the project description that you are interested in

The conditions for eligibility of home fees status are complex and you will need to seek advice if you have moved to or from the UK (or Republic of Ireland) within the past 3 years or have applied for settled status under the EU Settlement Scheme.


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 Friday 8 January 2021 2359 GMT .  Interviews will be held between 8th and 19th February 2021.  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:8th January 2021
Value:£15,285 per annum for 2020-21
Duration of award:per year
Contact: PGR Enquiries