Warming experiments are increasingly relied on to estimate plant responses to global climate change. For experiments to provide meaningful predictions of future responses, they should reflect the empirical record of responses to temperature variability and recent warming, including advances in the timing of flowering and leafing. We compared phenology (the timing of recurring life history events) in observational studies and warming experiments spanning four continents and 1,634 plant species using a common measure of temperature sensitivity (change in days per degree Celsius). We show that warming experiments underpredict advances in the timing of flowering and leafing by 8.5-fold and 4.0-fold, respectively, compared with long-term observations. For species that were common to both study types, the experimental results did not match the observational data in sign or magnitude. The observational data also showed that species that flower earliest in the spring have the highest temperature sensitivities, but this trend was not reflected in the experimental data. These significant mismatches seem to be unrelated to the study length or to the degree of manipulated warming in experiments. The discrepancy between experiments and observations, however, could arise from complex interactions among multiple drivers in the observational data, or it could arise from remediable artefacts in the experiments that result in lower irradiance and drier soils, thus dampening the phenological responses to manipulated warming. Our results introduce uncertainty into ecosystem models that are informed solely by experiments and suggest that responses to climate change that are predicted using such models should be re-evaluated.

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Forecasting how species and ecosystems will respond to climate change has been a major aim of ecology in recent years. Much of this research has focused on phenology -the timing of life-history events. Phenology has well-demonstrated links to climate, from genetic to landscape scales; yet our ability to explain and predict variation in phenology across species, habitats and time remains poor. Here, we outline how merging approaches from ecology, climate science and evolutionary biology can advance research on phenological responses to climate variability. Using insight into seasonal and interannual climate variability combined with niche theory and community phylogenetics, we develop a predictive approach for species' reponses to changing climate. Our approach predicts that species occupying higher latitudes or the early growing season should be most sensitive to climate and have the most phylogenetically conserved phenologies. We further predict that temperate species will respond to climate change by shifting in time, while tropical species will respond by shifting space, or by evolving. Although we focus here on plant phenology, our approach is broadly applicable to ecological research of plant responses to climate variability.

Analyses of datasets throughout the temperate midlatitude regions show a widespread tendency for species to advance their springtime phenology, consistent with warming trends over the past 20-50 y. Within these general trends toward earlier spring, however, are species that either have insignificant trends or have delayed their timing. Various explanations have been offered to explain this apparent nonresponsiveness to warming, including the influence of other abiotic cues (e.g., photoperiod) or reductions in fall/winter chilling (vernalization). Few studies, however, have explicitly attributed the historical trends of nonresponding species to any specific factor. Here, we analyzed long-term data on phenology and seasonal temperatures from 490 species on two continents and demonstrate that (i) apparent nonresponders are indeed responding to warming, but their responses to fall/winter and spring warming are opposite in sign and of similar magnitude; (ii) observed trends in first flowering date depend strongly on the magnitude of a given species' response to fall/winter vs. spring warming; and (iii) inclusion of fall/winter temperature cues strongly improves hindcast model predictions of long-term flowering trends compared with models with spring warming only. With a few notable exceptions, climate change research has focused on the overall mean trend toward phenological advance, minimizing discussion of apparently nonresponding species. Our results illuminate an understudied source of complexity in wild species responses and support the need for models incorporating diverse environmental cues to improve predictability of community level responses to anthropogenic climate change.growing season | ecological forecasting W ithin general trends toward earlier spring (1-7), observed cases of species and ecosystems that have not advanced their phenology, or have even delayed it, appear paradoxical, especially when made in temperate regions experiencing significant warming (2)(3)(4)(5)(8)(9)(10). The typical interpretation of this pattern has been that nonresponders are relatively insensitive to spring warming, whereas species showing delays are often viewed as statistical noise or evidence for unknown confounding factors at play. However, physiological work on model species has shown that the timing of flowering is controlled by multiple, complex pathways related to temperature forcing at different times during the plant life cycle (11). One important pathway is vernalizationchilling requirements that must be met before a plant is able to respond to spring warming (12, 13). These experimental studies suggest that warm temperatures during the vernalization period (typically fall and winter) can delay dormancy or the fulfillment of chilling requirements (14), thereby delaying spring events, such as flowering (13-18). Modeling and laboratory studies further suggest that this effect could temper phenological advances projected from climate warming (19,20).The importance of vernalization for explaining negligible or delayed ...