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Alan Grafen FRS is a Scottish ethologist and evolutionary biologist. He currently teaches and undertakes research at St John's College, Oxford.[1] Along with regular contributions to scientific journals, Grafen is known publicly for his work as co-editor (with Mark Ridley) of the 2006 festschrift Richard Dawkins: How a Scientist Changed the Way We Think,[2] honouring the achievements of his colleague and former academic advisor. He has worked extensively in the field of biological game theory, and, in 1990, devised a model showing that Zahavi's well-known handicap principle could theoretically exist in natural populations.[3][4]

Alan Grafen
CitizenshipUnited Kingdom
Scientific career
FieldsEthology, Evolutionary biology
InstitutionsUniversity of Oxford
ThesisThe economics of evolutionary stability (1984)
Doctoral advisorRichard Dawkins
Doctoral studentsLaurence Hurst

He also published a seminal paper in the field of phylogenetic comparative methods, in which he demonstrated how the tools of generalized least squares could be applied to perform phylogenetically informed statistical analyses.[5]

Grafen was elected a Fellow of the Royal Society in 2011.[6]


  • Hails, Rosemary; Grafen, Alan (2002). Modern statistics for the life sciences. Oxford [Oxfordshire]: Oxford University Press. ISBN 0-19-925231-9.


  1. ^ Alan Grafen's Web Page at Oxford University
  2. ^ Ridley, Mark; Grafen, Alan (2006). Richard Dawkins: how a scientist changed the way we think: reflections by scientists, writers, and philosophers. Oxford [Oxfordshire]: Oxford University Press. ISBN 0-19-929116-0.
  3. ^ Grafen, A. (1990). "Biological signals as handicaps". Journal of Theoretical Biology. 144 (4): 517–546. doi:10.1016/S0022-5193(05)80088-8. PMID 2402153.
  4. ^ Grafen, Alan (1 June 1990). "Sexual selection unhandicapped by the Fisher process". Journal of Theoretical Biology. 144 (4): 473–516. doi:10.1016/s0022-5193(05)80087-6.
  5. ^ Grafen, A. 1989. The phylogenetic regression. Phil. Trans. Royal. Soc. Lond. B 326:119-157.
  6. ^ "Professor Alan Grafen FRS". Royal Society. Retrieved 18 March 2012.