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Bet-hedging—a strategy that reduces fitness variance at the expense of lower mean fitness among different generations—is thought to evolve as a biological adaptation to environmental unpredictability. Despite widespread use of the bet-hedging concept, most theoretical treatments have largely made unrealistic demographic assumptions, such as non-overlapping generations and fixed or infinite population sizes. Here, we extend the concept to consider overlapping generations by defining bet-hedging as a strategy with lower variance and mean per capita growth rate across different environments. We also define an opposing strategy—the rising-tide—that has higher mean but also higher variance in per capita growth. These alternative strategies lie along a continuum of biological adaptions to environmental fluctuation. Using stochastic Lotka–Volterra models to explore the evolution of the rising-tide versus bet-hedging strategies, we show that both the mean environmental conditions and the temporal scales of their fluctuations, as well as whether population dynamics are discrete or continuous, are crucial in shaping the type of strategy that evolves in fluctuating environments. Our model demonstrates that there are likely to be a wide range of ways that organisms with overlapping generations respond to environmental unpredictability beyond the classic bet-hedging concept.

The reproduction of the plateau pika (Ochotona curzoniae) was investigated in Guoluo District at an elevation of 4,000 m on the Qinghai-Tibetan plateau, China, from April 2007 to August 2008. Reproduction was seasonal, and the breeding season lasted from April to late June/early July. Adults produced two litters in each year, and the mean litter size, estimated by counting the number of embryos in utero, was 3.3±0.1 (n=52) in 2007 and 3.2±0.1 (n=66) in 2008. The timing of reproduction showed high inter-annual variation; lower precipitation and the concomitant delay in spring vegetation phenology may have retarded the onset of the breeding season in 2007 compared with 2008. The most frequent litter sizes were 3 and 4, which together comprised 71.2% and 83.3% of litters in May and June of 2007 and 2008, respectively.Compared with previous studies, reproduction was highly variable between geographic areas. Pikas produced between one and five litters per year in different regions of the plateau over different breeding seasons. This geographic and inter-annual variation appeared to be associated with the duration of plant growth at each site, suggesting that plateau pikas adjust the length of their breeding season to match the period when sufficient energy is available to support the high energy demands of reproduction.