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Department of Zoology



I am an evolutionary biologist who works with butterflies and other insects. I have always been fascinated by the evolution of butterfly wing patterns and, in particular, by their eye-like markings known as eyespots. Can we explain the evolution of the diversity of these patterns both in terms of how they function and are influenced by natural selection, and how they are painted on the wings during development of the butterfly in the caterpillar and pupa? The mechanisms of development and physiology may contribute, alongside natural selection, to the patterns of diversity we see occurring in evolution.

Our multidiscipinary research on an African butterfly, Bicyclus anynana, is enabling us not only to tackle the issues of why eyespots matter in butterfly ecology but also how they are made by genes that regulate specific developmental mechanisms. In turn we can begin to explore the extent to which the way in which development builds morphologies can contribute to shaping evolutionary trajectories, for example via different forms of bias or constraint. B. anynana inhabits a wet-dry seasonal environment and shows striking phenotypic plasticity or polyphenism; the detailed study of this phenomenon has led us to investigate the roles of suites of metabolic, physiological, morphological and life history traits (including rates of aging) in ecological adaptation. The wing androconia of males of B. anynana are also now known to produce a species-specific blend of male sex pheromones that are involved in mate choice and sexual selection.

A new ERC-funded project known as EMARES will use our knowledge from these studies of a single model species to establish surveys and further experimental work to explore the processes of ecological diversification and speciation among the 250 or so species of these butterflies in the Old World tropics.


Key publications: 

Brakefield, P.M., Gates, J., Keys, D., Kesbeke, F., Wijngaarden, P.J., Monteiro, A., French, V., and Carroll, S.B. (1996). Development, plasticity and evolution of butterfly eyespot patterns. Nature 384, 236-242.

Beldade, P., Koops, K., and Brakefield, P.M. (2002). Developmental constraints versus flexibility in morphological evolution. Nature 416, 844-847.

Brakefield, P.M., and Reitsma, N. (1991). Phenotypic Plasticity, Seasonal Climate and the Population Biology of Bicyclus Butterflies (Satyridae) in Malawi. Ecol. Entomol. 16, 291-303.

Saccheri, I.J., Brakefield, P.M., and Nichols, R.A. (1996). Severe inbreeding depression and rapid fitness rebound in the butterfly Bicyclus anynana (Satyridae). Evolution 50, 2000-2013.

Joron, M., and Brakefield, P.M. (2003). Captivity masks inbreeding effects on male mating success in butterflies. Nature 424, 191-194.

Other publications: 

Bacquet, P.M.B.*, Brattström, O.*, Wang, H-L., Allen, C.E., Löfstedt, C., Brakefield, P.M. & Nieberding, C.M. (2015). Selection on male sex pheromone composition contributes to butterfly reproductive isolation. Proceedings of the Royal Society B, 282: 20142734. (* co-first authors)

Oostra, V., Mateus, A.R.A., van der Burg, K.R.L., Piessens, T., van Eijk, M., Brakefield, P.M., Beldade, P., Zwaan, B.J. (2014). Ecdysteroid hormones link the juvenile environment to alternative adult life histories in a seasonal insect. The American Naturalist

van Bergen, E., Brakefield, P.M., Heuskin, S., Zwaan, B.J., and Nieberding, C.M. (2013). The scent of inbreeding: a male sex pheromone betrays inbred males. Proc. R. Soc. B-Biol. Sci. 280.

Oostra, V., de Jong, M.A., Invergo, B.M., Kesbeke, F., Wende, F., Brakefield, P.M., and Zwaan, B.J. (2011). Translating environmental gradients into discontinuous reaction norms via hormone signalling in a polyphenic butterfly. Proc. R. Soc. B-Biol. Sci. 278, 789-797.

Saastamoinen, M., van der Sterren, D., Vastenhout, N., Zwaan, B.J., and Brakefield, P.M. (2010). Predictive Adaptive Responses: Condition-Dependent Impact of Adult Nutrition and Flight in the Tropical Butterfly Bicyclus anynana. Am. Nat. 176, 686-698.

Saenko, S.V., Brakefield, P.M., and Beldade, P. (2010). Single locus affects embryonic segment polarity and multiple aspects of an adult evolutionary novelty. BMC Biology 8.

Lommen, S.T.E., Saenko, S.V., Tomoyasu, Y., and Brakefield, P.M. (2009). Development of a wingless morph in the ladybird beetle, Adalia bipunctata. Evol Dev 11, 278-289.

Professor of Zoology (Emeritus)

Contact Details

Museum Room G.16
01223 (3)36612 or (3)36623
Accepting applications for PhD students.