SummaryMast seeding is a crucial population process in many tree species, but its spatio-temporal patterns and drivers at the continental scale remain unknown .Using a large dataset (8000 masting observations across Europe for years 1950-2014) we analysed the spatial pattern of masting across the entire geographical range of European beech, how it is influenced by precipitation, temperature and drought, and the temporal and spatial stability of masting-weather correlations.Beech masting exhibited a general distance-dependent synchronicity and a pattern structured in three broad geographical groups consistent with continental climate regimes. Spearman's correlations and logistic regression revealed a general pattern of beech masting correlating negatively with temperature in the summer 2 yr before masting, and positively with summer temperature 1 yr before masting (i.e. 2T model). The temperature difference between the two previous summers (DeltaT model) was also a good predictor. Moving correlation analysis applied to the longest eight chronologies (74-114 yr) revealed stable correlations between temperature and masting, confirming consistency in weather cues across space and time.These results confirm widespread dependency of masting on temperature and lend robustness to the attempts to reconstruct and predict mast years using temperature data.

Highly variable and synchronised production of seeds by plant populations, known as masting, is implicated in many important ecological processes, but how it arises remains poorly understood. The lack of experimental studies prevents underlying mechanisms from being explicitly tested, and thereby precludes meaningful predictions on the consequences of changing environments for plant reproductive patterns and global vegetation dynamics. Here we review the most relevant proximate drivers of masting and outline a research agenda that takes the biology of masting from a largely observational field of ecology to one rooted in mechanistic understanding. We divide the experimental framework into three main processes: resource dynamics, pollen limitation and genetic and hormonal regulation, and illustrate how specific predictions about proximate mechanisms can be tested, highlighting the few successful experiments as examples. We envision that the experiments we outline will deliver new insights into how and why masting patterns might respond to a changing environment.

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Aim Climate limits the potential distribution ranges of species. Establishment and growth of individuals at range margins is assumed to be more limited by extreme events such as drought or frost events than in the centre of their range. We explore whether the growth of beech is more sensitive to drought towards the dry distribution margin and more sensitive to frost towards the cold distribution margin. Furthermore, we aim to gain insight into the adaptive potential of beech towards both the dry and cold distribution margins. Location European gradient from the dry (Spain) to the cold (Poland, Sweden) distribution margin of beech. Taxon European beech (Fagus sylvatica L.). Methods We applied a range‐wide dendroecological study to analyse spatial and temporal trends in climate–growth relationships. We further investigated negative growth anomalies and growth synchrony towards the range margins. Results We found beech to be drought sensitive across its whole range, except at the dry distribution margin. Furthermore, sensitivity to winter temperature was not found in the centre or at the cold distribution margin, but at the southern distribution margin. Growth synchrony was lower at the dry than at the cold distribution margin. Main conclusions Beech seems to be adapted to drought at the dry distribution margin with a high adaptive potential indicated by the lowest growth synchrony along the gradient. At the cold distribution margin, cold events in winter and spring were less important for growth than drought. Still, the importance of spring frost for beech growth appears to increase in recent decades. Considering a projected north‐eastward shift of the distribution range, beech is likely facing drought stress in combination with spring frost risk at the cold margin which could lead to a hampered range expansion.

Climate change is altering patterns of seed production worldwide with consequences for population recruitment and migration potential. For the many species that regenerate through synchronized, quasiperiodic reproductive events termed masting, these changes include decreases in the synchrony and interannual variation in seed production. This breakdown in the occurrence of masting features harms reproduction by decreasing the efficiency of pollination and increasing seed predation. Changes in masting are often paralleled by warming temperatures, but the underlying proximate mechanisms are unknown. We used a unique 39‐year study of 139 European beech (Fagus sylvatica) trees that experienced masting breakdown to track the seed developmental cycle and pinpoint phases where weather effects on seed production have changed over time. A cold followed by warm summer led to large coordinated flowering efforts among plants. However, trees failed to respond to the weather signal as summers warmed and the frequency of reproductive cues changed fivefold. Less synchronous flowering resulted in less efficient pollination that further decreased the synchrony of seed maturation. As global temperatures are expected to increase this century, perennial plants that fine‐tune their reproductive schedules based on temperature cues may suffer regeneration failures.

Spatial synchrony is the tendency of spatially separated populations to display similar temporal fluctuations. Synchrony affects regional ecosystem functioning, but it remains difficult to disentangle its underlying mechanisms. We leveraged regression on distance matrices and geography of synchrony to understand the processes driving synchrony of European beech masting over the European continent. Masting in beech shows distance‐decay, but significant synchrony is maintained at spatial scales of up to 1,500 km. The spatial synchrony of the weather cues that drive interannual variation in reproduction also explains the regional spatial synchrony of masting. Proximity played no apparent role in influencing beech masting synchrony after controlling for synchrony in environmental variation. Synchrony of beech reproduction shows a clear biogeographical pattern, decreasing from the northwest to southeast Europe. Synchrony networks for weather cues resemble networks for beech masting, indicating that the geographical structure of weather synchrony underlies the biogeography of masting synchrony. Our results support the hypothesis that environmental factors, the Moran effect, are key drivers of spatial synchrony in beech seed production at regional scales. The geographical patterns of regional synchronization of masting have implications for regional forest production, gene flow, carbon cycling, disease dynamics, biodiversity, and conservation.

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