Evolutionary Ecology of Senescence and a Reassessment of Williams' 'Extrinsic Mortality' Hypothesis.

The evolutionary theory of senescence underpins research in life history evolution and the biology of aging. In 1957 G.C. Williams predicted that higher adult death rates select for earlier senescence and shorter length of life, but preadult mortality does not matter to the evolution of senescence. This was subsequently interpreted as predicting that senescence should be caused by 'extrinsic' sources of mortality. This idea still motivates empirical studies, although formal, mathematical theory shows it is wrong. It has nonetheless prospered because it offers an intuitive explanation for patterns observed in nature. We review the flaws in Williams' model, explore alternative explanations for comparative patterns that are consistent with the evolutionary theory of senescence, and discuss how hypotheses based on it can be tested. We argue that focusing on how sources of mortality affect ages differently offers greater insight into evolutionary processes.