Telephone number: 01865 (2)77438

Social media: @GrafenAlan

Role: College Lecturer (Fellow of St John's College)

Prof Alan Grafen FRS


I have an undergraduate degree in Experimental Psychology, a masters degree in Economics (supervisor Sir James (Jim) Mirrlees) and a doctorate in Zoology (supervisor Richard Dawkins). I apply in biology, to evolutionary and statistical topics, the modelling skills I learnt as an economist. Currently an Official Fellow of St John’s College, and Honorary Fellow of Jesus College. Elected to the Royal Society in 2011.

Research interests

My research involves applying mathematical and logical theorising to evolutionary problems. I have worked on animal signals, sexual selection, kin selection, kin recognition, the statistics of cross-species comparisons and optimality models of sequences of behaviour. Since 2000, I have been constructing a mathematical framework called Formal Darwinism that aims to link Darwinian ideas of biological design with population genetics. Most recently, I have been incorporating Fisher’s Fundamental Theorem of Natural Selection into the project.


I teach courses that have mathematical content, as well as those close to Evolution and Animal Behaviour. I give tutorials in the Ecology and Evolution course to first years, in the Evolution, Animal Behaviour and Quantitative Methods courses to second years, and in Social Evolution to third years.


Biological Sciences

Selected publications

  • A. Grafen. 2015. Biological fitness and the Fundamental Theorem of Natural Selection. American Naturalist 186, 1-14. doi: 10.1086/681585.
  • Biernaskie, J.M., Grafen, A. and J.C. Perry. 2014. The evolution of index signals to avoid the cost of dishonesty. Proc. R. Roy. Soc. B 281, 20140876. doi: 10.1098/rspb.2014.0876.
  • Batty, C.J.K., Crewe, P., Grafen, A. and R. Gratwick. 2014. Foundations of a mathematical theory of darwinism. Journal of mathematical Biology 69, 295-334. doi: 10.1007/s00285-013-0706-2.
  • Grafen, A. 2014. The formal darwinism project in outline. Biology and Philosophy, 29, 155-174. doi: 10.1007/s10539-013-9414-y.