Reproductive synchrony or desynchrony of primate females influences number and fitness of males in mating systems. Langur monkey populations provide a natural experiment for observing alternative female strategies of confusing or concentrating paternity. Where females escape seasonal reproductive constraints, they desynchronize fertility and show visible cues (menstruation), enabling single males to monopolize matings. This increases female fitness by reducing food competition. Where langurs are seasonally constrained, females conceal fertility, confusing paternity and reducing infanticide. These case studies illuminate how hominin females could increase male numbers and investment. Fitness payoffs to male investors will be affected by degree of reproductive seasonal constraint, and by females either concealing or confusing menstrual cues of imminent fertility. Among ancestors of modern humans and Neanderthals these strategies diverged. Under pressure of encephalization, modern human female ancestors, less seasonally constrained, pursued a strategy of cosmeticization of menstrual signals. This Female Cosmetic Coalitions model accounts for the African Middle Stone Age record of pigment use. Among Neanderthals, strategies alternated. Severe seasonality during glacial cycles tied Neanderthal males into pair-bonds, suppressing cosmetic signaling. Only during interglacials when seasonality relaxed would Neanderthal females require blood-red cosmetics. Our Seasonality Thermostat model explains why European ochre use correlates with climate through the Middle to Late Pleistocene.
INTRODUCTION
There is a growing consensus that evolving Homo mothers relied on cooperative breeding to raise larger-bodied, larger-brained offspring (e.g., Aiello and Key 2002;Barrickman et al. 2008;Burkart et al. 2009;Robson and Wood 2008;Shultz et al. 2012;van Schaik et al. 2006 Models for cultural levels of human cooperation (Boyd and Richerson 1992, 2005Gintis et al. 2003;Henrich and Boyd 2001;Henrich et al. 2003Henrich et al. , 2006Nowak and Sigmund 2005;Richerson et al. 2003), invoke multilevel and group selection processes, but ignore what is most difficult to explain-how to secure cooperation between the sexes, given the trade-off between investing in current offspring versus future mating opportunities (cf. Bowles 2006). One game theory model (Key 1998(Key , 2000 incorporated the differences of reproductive energetic costs to specify conditions for the evolution of cooperation that can be practically applied to the fossil record. Yet even these conditions do not identify how females induced males to subsidize their reproduction (Kaplan et al. 2000;Marlowe 2001). Recent models (Gavrilets 2012) highlight the difficulty of shifting from systems of significant male mating competition into productive male investment, where males face a social dilemma of ceding fitness advantage to free-riding rivals.If we can define the mechanisms available to Homo females giving them leverage against male defection through the Early to Middle Plei...