Sex ratio bias and reproductive strategies: what sex to produce when?

Several theories predict the evolution of bias in progeny sex ratio based on variations in maternal or offspring reproductive value. For mammals, however, tests of sex-bias theories have produced inconsistent results, and no clear patterns have emerged. Each theory is based on assumptions that are difficult to satisfy, and empirical tests require large data sets. Using a long-term study on bighorn sheep (Ovis canadensis), we identified several parameters that influence progeny sex ratio according to maternal state. For older females, progeny sex ratio was affected by an interaction between reproductive strategy and environmental conditions. When conditions were good, old females reproduced every year but minimized fitness costs by producing daughters. When conditions were poor, old females produced more sons but did not reproduce every year. Sons of older females were of similar mass to those born to younger females under poor conditions but were smaller and likely disadvantaged under good environmental conditions. For young and prime-aged females, progeny sex ratio was independent of environmental conditions. Environmental conditions and age should be considered when studying sex ratio bias, which appears to be a function of maternal state rather than of maternal condition. We suggest that a conservative reproductive strategy drives progeny sex ratio in older females according to the "cost of reproduction hypothesis." By manipulating offspring sex ratio, older females reduced the cost of reproduction and increased their expected fitness returns.