Is biasing offspring sex ratio adaptive? a test of Fisher’s principle across multiple generations of a wild mammal in a fluctuating environment

Fisher’s principle explains that population sex ratio in sexually reproducing organisms is maintained at 1:1 due to negative frequency-dependent selection, such that individuals of the rare sex realize greater reproductive opportunity than individuals of the more common sex until equilibrium is reached. If biasing offspring sex ratio towards the rare sex is adaptive, individuals that do so should have a higher number of grandoffspring. In a wild population of North American red squirrels (Tamiasciurus hudsonicus) that experiences fluctuations in resource abundance and population density, we show that overall across 26 years, the secondary sex ratio was 1:1; however, stretches of years during which adult sex ratio was biased did not yield offspring sex ratios biased towards the rare sex. Females that had litters biased towards the rare sex did not have more grandoffspring. Critically, adult sex ratio was not temporally autocorrelated across years, thus the population sex ratio experience...