In 1957, George Williams [ 1. Williams G.C. Pleiotropy, natural selection, and the evolution of senescence. Evolution. 1957; 11: 398-411 Crossref Google Scholar ] argued that higher, age-independent, adult mortality should be correlated with more rapid senescence. Charlesworth [ 2. Charlesworth B. Evolution in Age-Structured Populations. Cambridge University Press, 1980 Google Scholar ] and most subsequent studies have added the adjective 'extrinsic' to mean that the age-independent mortality was due to sources 'external' to the organism. We now know that neither age- nor condition-independent increases in extrinsic mortality are necessary for more rapid senescence to evolve, and analyses have also shown that higher mortality can lead to the evolution of slower senescence [ 3. Abrams P.A. Does increased mortality favor the evolution of more rapid senescence?. Evolution. 1993; 47: 877-887 Crossref PubMed Google Scholar , 4. Williams P.D. Day T. Antagonistic pleiotropy, mortality source interactions, and the evolutionary theory of senescence. Evolution. 2003; 57: 1478-1488 Crossref PubMed Scopus (198) Google Scholar ]. Indeed, there are empirical examples where an added source of mortality seems to have extended lifespan [ 5. Reznick D.N. et al. The effect of extrinsic mortality on the evolution of senescence in guppies. Nature. 2004; 431: 1095-1099 Crossref PubMed Scopus (334) Google Scholar ]. Nevertheless, Williams' hypothesis remains popular [ 6. Gaillard J.-M. Lemaitre J.-F. The Williams' legacy: a critical reappraisal of his nine predictions about the evolution of senescence. Evolution. 2017; 71: 2768-2785 Crossref PubMed Scopus (68) Google Scholar ].
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