 Along with regular contributions to scientific journals, Grafen is known publicly for his work as co-editor (with Mark Ridley) of the 2006 festschrift "", 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.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.