What lies beneath? Population dynamics conceal pace-of-life and sex ratio variation, with implications for resilience to environmental change.

Life-history and pace-of-life syndrome theory predict that populations are comprised of individuals exhibiting different reproductive schedules and associated behavioural and physiological traits, optimized to prevailing social and environmental factors. Changing weather and social conditions provide in situ cues altering this life-history optimality; nevertheless, few studies have considered how tactical, sex-specific plasticity over an individual's lifespan varies in wild populations and influences population resilience. We examined the drivers of individual life-history schedules using 31 years of trapping data and 28 years of pedigree for the European badger (Meles meles L.), a long-lived, iteroparous, polygynandrous mammal that exhibits heterochrony in the timing of endocrinological puberty in male cubs. Our top model for the effects of environmental (social and weather) conditions during a badger's first year on pace-of-life explained <10% of variance in the ratio of fertility to age at first reproduction (F/α) and lifetime reproductive success. Conversely, sex ratio (SR) and sex-specific density explained 52.8% (males) and 91.0% (females) of variance in adult F/α ratios relative to the long-term population median F/α. Weather primarily affected the sexes at different life-history stages, with energy constraints limiting the onset of male reproduction but playing a large role in female strategic energy allocation, particularly in relation to ongoing mean temperature increases. Furthermore, the effects of social factors on age of first reproduction and year-to-year reproductive success covaried differently with sex, likely due to sex-specific responses to potential mate availability. For females, low same-sex densities favoured early primiparity; for males, instead, up to 10% of yearlings successfully mated at high same-sex densities. We observed substantial SR dynamism relating to differential mortality of life-history strategists within the population, and propose that shifting ratios of 'fast' and 'slow' life-history strategists contribute substantially to population dynamics and resilience to changing conditions.