| Citation | Alan Grafen is distinguished for mathematical and conceptual contributions to behavioural ecology. He introduced 'split sex ratios', now a quantitatively exact area of empirical research. He successfully modelled Zahavi s handicap principle, allowing a new and productive understanding of how signals can function in nature. Grafen's style of modelling represents organisms as sophisticated actors, allowing the hypotheses and intuitions of empirical biologists to be represented in precise, mathematical form, including general foundational questions about fitness and its optimisation. A rederivation of inclusive fitness, and a general construction of Fisher s reproductive value, are recent examples. Advanced abstract mathematics is used where required. |