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He took an early interest in lepidoptera and ecological genetics following the work of E.B. Ford. Later he moved into studying ladybirds.
His website defines his "keyword" subject areas as:
Evolutionary genetics, ecological genetics, Insecta, polymorphism, melanism, Peppered moth, male-killers, Wolbachia, sexually transmitted disease, reproductive costs, reproductive strategies, ultra-violet patterns in insects, sexual selection, Lepidoptera, Ladybird beetles (Coccinellidae).
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