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Theory of social evolution: cooperation and conflict at multiple levels of biological organisation

School of Biology

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Prof A Gardner , Prof G Ruxton No more applications being accepted Funded PhD Project (European/UK Students Only)

About the Project

Natural selection explains the appearance of design in the living world. But at what level is this design expected to manifest – gene, individual, society – and what is its function? Social evolution provides a window on this problem, by pitting the interests of genes, individuals and societies against each other.

I invite applications for a PhD studentship in my research group at the School of Biology, University of St Andrews, UK. I’m looking for a biology graduate who has a strong interest in social evolution theory, or an economics / mathematics / philosophy / physics graduate with a strong interest in social behaviour.

Current research in my lab involves development of general theory – using kin selection, multilevel selection, game theory and theoretical population genetics approaches – and application of mathematical and simulation models that are tailored to the biology of real organisms, from microbes to insects to humans (see for more details). To this end, I strongly encourage interactions within the wider grouping of theoretical and empirical biologists in St Andrews, as well as with collaborators further afield.

If social evolution really fascinates you, and you are a careful thinker, then you will flourish in the type of project that I enjoy supervising.
Please direct informal enquiries to Prof Andy Gardner ([Email Address Removed]).

Funding Notes

Eligibility requirements: Upper second-class degree in Biology or a related area.

This PhD will provide 3.5 years of ERC funding for UK or EU students, including fees, stipend and research and travel expenses.


1. Micheletti AJC, Ruxton GD & Gardner A. 2018. Why war is a man’s game. Proceedings of the Royal Society of London Series B — Biological Sciences 285: 20180975.
2. Gardner A & Úbeda F. 2017. The meaning of intragenomic conflict. Nature Ecology & Evolution 1: 1807.
3. Faria GS, Varela SAM & Gardner A. 2017. Sexual selection modulates genetic conflicts and patterns of genomic imprinting. Evolution 71: 526.
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