Summary 1.Variation in climate is predicted to increase over much of the planet this century. Forecasting species persistence with climate change thus requires understanding of how populations respond to climate variability, and the mechanisms underlying this response. Variable rainfall is well known to drive fluctuations in annual plant populations, yet the degree to which population response is driven by between-year variation in germination cueing, water limitation or competitive suppression is poorly understood. 2. We used demographic monitoring and population models to examine how three seed banking, rare annual plants of the California Channel Islands respond to natural variation in precipitation and their competitive environments. Island plants are particularly threatened by climate change because their current ranges are unlikely to overlap regions that are climatically favourable in the future. 3. Species showed 9 to 100-fold between-year variation in plant density over the 5-12 years of censusing, including a severe drought and a wet El Niño year. During the drought, population sizes were low for all species. However, even in non-drought years, population sizes and per capita growth rates showed considerable temporal variation, variation that was uncorrelated with total rainfall. These population fluctuations were instead correlated with the temperature after the first major storm event of the season, a germination cue for annual plants. 4. Temporal variation in the density of the focal species was uncorrelated with the total vegetative cover in the surrounding community, suggesting that variation in competitive environments does not strongly determine population fluctuations. At the same time, the uncorrelated responses of the focal species and their competitors to environmental variation may favour persistence via the storage effect. 5. Population growth rate analyses suggested differential endangerment of the focal annuals. Elasticity analyses and life table response experiments indicated that variation in germination has the same potential as the seeds produced per germinant to drive variation in population growth rates, but only the former was clearly related to rainfall. 6. Synthesis . Our work suggests that future changes in the timing and temperatures associated with the first major rains, acting through germination, may more strongly affect population persistence than changes in season-long rainfall.

A major challenge in forecasting the ecological consequences of climate change is understanding the relative importance of changes to mean conditions vs. changes to discrete climatic events, such as storms, frosts, or droughts. Here we show that the first major storm of the growing season strongly influences the population dynamics of three rare and endangered annual plant species in a coastal California (USA) ecosystem. In a field experiment we used moisture barriers and water addition to manipulate the timing and temperature associated with first major rains of the season. The three focal species showed two- to fivefold variation in per capita population growth rates between the different storm treatments, comparable to variation found in a prior experiment imposing eightfold differences in season-long precipitation. Variation in germination was a major demographic driver of how two of three species responded to the first rains. For one of these species, the timing of the storm was the most critical determinant of its germination, while the other showed enhanced germination with colder storm temperatures. The role of temperature was further supported by laboratory trials showing enhanced germination in cooler treatments. Our work suggests that, because of species-specific cues for demographic transitions such as germination, changes to discrete climate events may be as, if not more, important than changes to season-long variables.

Ecologists increasingly suspect that climate change will directly impact species physiology, demography, and phenology, but also indirectly affect these measures via changes to the surrounding community. Unfortunately, few studies examine both the direct and indirect pathways of impact. Doing so is important because altered competitive pressures can reduce or magnify the direct responses of a focal species to climate change. Here, we examine the effects of changing rainfall on three rare annual plant species in the presence and absence of competition on the California Channel Islands. We used rain-out shelters and hand watering to exclude and augment early, late, and season-long rainfall, spanning the wide range of precipitation change forecast for the region. In the absence of competition, droughts reduced the population growth rates of two of three focal annuals, while increased rainfall was only sometimes beneficial. As compared to the focal species, the dominant competitors were more sensitive to the precipitation treatments, benefiting from increased season-long precipitation and harmed by droughts. Importantly, the response of two of three competitors to the precipitation treatments tended to be positively correlated with those of the focal annuals. Although this leads to the expectation that increased competition will counter the direct benefits of favorable conditions, such indirect effects of precipitation change proved weak to nonexistent in our experiment. Competitors had little influence on the precipitation response of two focal species, due to their low sensitivity to competition and highly variable precipitation responses. Competition did affect how our third focal species responded to precipitation change, but this effect only approached significance, and whether it truly resulted from competitor response to precipitation change was unclear. Our work suggests that even when competitors respond to climate change, these responses may have little effect on the focal species. Ultimately, the strength of the indirect effect depends on how strongly climate change alters competition, and how sensitive focal species are to changes in competition.

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