33Natural populations are exposed to seasonal variation in environmental factors that 34 simultaneously affect several demographic rates (survival, development, reproduction). The 35 resulting covariation in these rates determines population dynamics, but accounting for its 36 numerous biotic and abiotic drivers is a significant challenge. Here, we use a factor-analytic 37 approach to capture partially unobserved drivers of seasonal population dynamics. We use 40 38 years of individual-based demography from yellow-bellied marmots (Marmota flaviventer) to fit 39 and project population models that account for seasonal demographic covariation using a latent 40 variable. We show that this latent variable, by producing positive covariation among winter 41 demographic rates, depicts a measure of environmental quality. Simultaneous, negative 42 responses of winter survival and reproductive-status change to declining environmental quality 43 result in a higher risk of population quasi-extinction, regardless of summer demography where 44 recruitment takes place. We demonstrate how complex environmental processes can be 45 summarized to understand population persistence in seasonal environments. 47 Effects of environmental change on survival, growth, and reproduction are typically investigated 48 based on annual transitions among life-history stages in structured population models (Salguero-49 GĂłmez et al., 2016; Paniw et al., 2018). However, all natural ecosystems show some level of 50 seasonal fluctuations in environmental conditions, and numerous species have evolved life cycles 51 that are cued to such seasonality (Ruf et al., 2012; Varpe, 2017). For example, most temperate-52 and many arid-environment species show strong differences in survival and growth among 53 seasons, with reproduction being confined mostly to one season (Childs et al., 2011; Rushing et 54 al., 2017; Woodroffe et al., 2017). Species with highly adapted, seasonal life cycles are likely to 55 be particularly vulnerable to environmental change, even if they are relatively long-lived 56 (Jenouvrier et al., 2012; Campos et al., 2017; Paniw et al., 2019). This is because adverse 57 environmental conditions in the non-reproductive season may carry-over and negate positive 58 environmental effects in the reproductive season in which key life-history events occur (Marra et 59 al., 2015). For instance, in species where individual traits such as body mass determine 60 demographic rates, environment-driven changes in the trait distribution in one season can affect 61 trait-dependent demographic rates in the next season (Bassar et al., 2016; Paniw et al., 2019).
62Investigating annual dynamics, averaged over multiple seasons, may, therefore, obscure the 63 mechanisms that allow populations to persist under environmental change.
64Despite the potential to gain a more mechanistic view of population dynamics, modeling 65 the effects of seasonal environmental change is an analytically complex and data-hungry 66 endeavor (Benton et al., 2006; Bassar et al...