Growing up to help: understanding the developmental plasticity of social behaviour in cooperative societies. NERC GW4+, PhD In Mathematics studentship. Ref: 3345

About the award


Lead Supervisor

Prof Stuart Townley Department of Mathematics, College of Engineering, Mathematics and Physical Sciences, University of Exeter.

Additional Supervisors

Dr Sinead English School of Biological Sciences, University of Bristol.

Dr Bram Kuijper, Department of Biological Sciences, College of Life and Environmental Sciences, University of Exeter

Prof Jeremy Field  Department of Biological Sciences, College of Life and Environmental Sciences, University of Exeter.

Location: University of Exeter, Penryn Campus, Penryn, Cornwall TR10 9FE 

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 index-linked stipend for 3.5 years (currently £14,777 p.a. for 2018/19);
  • Payment of university tuition fees;
  • A research budget of £11,000 for an international conference, lab, field and research expenses;
  • A training budget of £4,000 for specialist training courses and expenses.

Up to 30 fully-funded studentships will be available across the partnership.

Students from EU countries who do not meet the residency requirements may still be eligible for a fees-only award but no stipend.  Applicants who are classed as International for tuition fee purposes are not eligible for funding.

Project details

Cooperatively breeding animals exhibit a broad variety of developmental trajectories, ranging from eusocial insects with irreversible specialization as queens versus workers, to vertebrates where individuals flexibly revert from helping to reproduction. Between those extremes lies a continuum of variation in social flexibility. Making sense of this variation is essential to understand the developmental basis of cooperation, yet testable predictions are currently lacking1,2. The current project aims to fill this gap by building evolutionary models in which we track the development of social behaviours in response to the environmental and social cues that individuals experience in early life (e.g. the amount of help received, variation in resources, presence of social parasites, weather). We will test the models using taxa such as sweat bees (Halictus) and silk wasps (Microstigmus) in which some populations contain eusocial nests while other populations of the same species are entirely non-social. The result should be a step change in our understanding of what drives the development of social behaviours, and how such behaviour will alter under climate change3.

Project Aims and Methods

1. Use kin selection theory to investigate the evolution of developmental plasticity in social traits Use a combination of simulation and analytical techniques to model a society of cooperative individuals. We then assess how variation in their abiotic environment (e.g. variation in the amounts of resources available) as well as their social environment (e.g. variation in the local numbers of males vs females, or random variation in helper numbers) affects the propensity of individuals to develop as helpers vs reproductives.

2. Correlate environmental variation with social phenotype in sweat bees and silk wasps Use data loggers to measure microclimate at field sites and assess how environmental variation (e.g. autocorrelations in rainfall) relates to social phenotype (e.g. helper number, division of reproduction among group-members).

3. Perform a meta-analysis relating cooperative vs uncooperative phenotypes to environmental variation Compile a database which associates environmental variation with social phenotype across cooperative taxa and solitary sister species.

4. Carry out manipulative field experiments to test model predictions. Experiments would be carried out at established field sites in the UK (sweat bees)4 and/or Ecuador (silk wasps), in which either the abiotic or the social environment will be manipulated.
The balance between aims 1-4 will be determined as the project and the interests of the student develop.


The student will be trained in:

Planning and conducting research from idea through to publication;

Developing evolutionary models of cooperative societies;

Data analysis and field techniques in South America and/or the UK;

Meta-analysis and individual-based simulations in C++

The student will benefit from the input of a team of supervisors with diverse expertise and across different disciplines, including mathematical modelling of biological systems (Townley, Kuijper [Exeter, UK]), field ecology (Field [Exeter UK], English [Bristol, UK]) evolutionary theory (Kuijper) and data analysis (Kuijper, English).


Fig.1 The sweat bee Halictus rubicundus, a socially polymorphic species. Dependent on environmental/social stimuli, females either nest solitarily or form social groups. NERC

Fig.2 A nest of a silk wasp of the neotropical genus Microstigmus. Silk wasp species are highly socially diverse, varying in (among others) division of labour and mating skew.

References / Background reading list

1.English, S., Browning, L. E. & Raihani, N. J. Developmental plasticity and social specialization in cooperative societies. Anim. Behav. 106, 37–42 (2015).

2.Kasper, C. et al. Genetics and developmental biology of cooperation. Mol. Ecol. 26, 4364–4377 (2017).

3.Schürch, R., Accleton, C. & Field, J. Consequences of a warming climate for social organisation in sweat bees. Behav. Ecol. Sociobiol. 70, 1131–1139 (2016).

4.Field, J., Paxton, R. J., Soro, A. & Bridge, C. Cryptic plasticity underlies a major evolutionary transition. Curr. Biol. 20, 2028–2031 (2010).

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.

Candidate Requirements

The candidate should have a strong interest in the major questions in ecology and evolution, such as why individuals perform cooperative behaviours as opposed to selfishness, or how collectives of individuals adapt to evolutionary change. The candidate should also have an interest in learning more about mathematical models in ecology & evolution, and should have experience of statistics and/or coding in a programming language like R.

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.
  • Two References (applicants are recommended to have a third academic referee, if the two academic referees are within the same department/school).

Reference information
You will be asked to name two referees as part of the application process.  It is your responsibility to ensure that your two referees email their references to, as we will not make requests for references directly; you must arrange for them to be submitted by 7 January 2019

References should be submitted 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 midnight on 7 January 2019.  Interviews will be held between 4 and 15 February 2019.

If you have any general enquiries about the application process please email  Project-specific queries should be directed to the 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:7th January 2019
Value:£14,777 per annum for 2018-19
Duration of award:per year
Contact: PGR Enquiries