Fifty years ago, Ernst Mayr published a hugely influential paper on the nature of causation in biology, in which he distinguished between proximate and ultimate causes. Mayr equated proximate causation with immediate factors (for example, physiology) and ultimate causation with evolutionary explanations (for example, natural selection). He argued that proximate and ultimate causes addressed different questions and were not alternatives. Mayr's account of causation remains widely accepted today, with both positive and negative ramifications. Several current debates in biology (for example, over evolution and development, niche construction, cooperation, and the evolution of language) are linked by a common axis of acceptance/rejection of May...