Why does women's fertility end in mid-life? Grandmothering and age at last birth.

Great apes, the other living members of our hominid family, become decrepit before the age of forty and rarely outlive their fertile years. In contrast, women - even in high mortality hunter-gatherer populations - usually remain healthy and productive well beyond menopause. The grandmother hypothesis aims to account for the evolution of this distinctive feature of human life history. Our previous mathematical simulations of that hypothesis fixed the end of female fertility at the age of 45, based on the similarities among living hominids, and then modeled the evolution of human-like longevity from an ancestral state, like that of the great apes, due only to grandmother effects. A major modification here allows the age female fertility ends to vary as well, directly addressing a version of the question, influentially posed by GC Williams six decades ago: Why isn't menopause later in humans? Our model is an agent-based model (ABM) that accounts for the coevolution of both expected adult lifespan and end of female fertility as selection maximizes reproductive value. We find that grandmother effects not only drive the population from an equilibrium representing a great ape-like longevity to a new human-like longevity, they also maintain the observed termination of women's fertility before the age of 50.

Grandmothering life histories and human pair bonding.

The evolution of distinctively human life history and social organization is generally attributed to paternal provisioning based on pair bonds. Here we develop an alternative argument that connects the evolution of human pair bonds to the male-biased mating sex ratios that accompanied the evolution of human life history. We simulate an agent-based model of the grandmother hypothesis, compare simulated sex ratios to data on great apes and human hunter-gatherers, and note associations between a preponderance of males and mate guarding across taxa. Then we explore a recent model that highlights the importance of mating sex ratios for differences between birds and mammals and conclude that lessons for human evolution cannot ignore mammalian reproductive constraints. In contradiction to our claim that male-biased sex ratios are characteristically human, female-biased ratios are reported in some populations. We consider the likelihood that fertile men are undercounted and conclude that the mate-guarding hypothesis for human pair bonds gains strength from explicit links with our grandmothering life history.

Grandmothering drives the evolution of longevity in a probabilistic model.

We present a mathematical model based on the Grandmother Hypothesis to simulate how human post-menopausal longevity could have evolved as ancestral grandmothers began to assist the reproductive success of younger females by provisioning grandchildren. Grandmothers׳ help would allow mothers to give birth to subsequent offspring sooner without risking the survival of existing offspring. Our model is an agent-based model (ABM), in which the population evolves according to probabilistic rules governing interactions among individuals. The model is formulated according to the Gillespie algorithm of determining the times to next events. Grandmother effects drive the population from an equilibrium representing a great-ape-like average adult lifespan in the lower twenties to a new equilibrium with a human-like average adult lifespan in the lower forties. The stochasticity of the ABM allows the possible coexistence of two locally-stable equilibria, corresponding to great-ape-like and human-like lifespans. Populations with grandmothering that escape the ancestral condition then shift to human-like lifespan, but the transition takes longer than previous models (Kim et al., 2012). Our simulations are consistent with the possibility that distinctive longevity is a feature of genus Homo that long antedated the appearance of our species.