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Extreme droughts generally decrease productivity in grassland ecosystems1,2,3 with negative consequences for nature’s contribution to people4,5,6,7. The extent to which this negative effect varies among grassland types and over time in response to multi-year extreme drought remains unclear. Here, using a coordinated distributed experiment that simulated four years of growing-season drought (around 66% rainfall reduction), we compared drought sensitivity within and among six representative grasslands spanning broad precipitation gradients in each of Eurasia and North America—two of the Northern Hemisphere’s largest grass-dominated regions. Aboveground plant production declined substantially with drought in the Eurasian grasslands and the effects accumulated over time, while the declines were less severe and more muted over time in the North American grasslands. Drought effects on species richness shifted from positive to negative in Eurasia, but from negative to positive in North America over time. The differing responses of plant production in these grasslands were accompanied by less common (subordinate) plant species declining in Eurasian grasslands but increasing in North American grasslands. Our findings demonstrate the high production sensitivity of Eurasian compared with North American grasslands to extreme drought (43.6% versus 25.2% reduction), and the key role of subordinate species in determining impacts of extreme drought on grassland productivity.more » « lessFree, publicly-accessible full text available January 29, 2026
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Abstract The relationship between stomatal traits and environmental drivers across plant communities has important implications for ecosystem carbon and water fluxes, but it has remained unclear. Here, we measure the stomatal morphology of 4492 species-site combinations in 340 vegetation plots across China and calculate their community-weighted values for mean, variance, skewness, and kurtosis. We demonstrate a trade-off between stomatal density and size at the community level. The community-weighted mean and variance of stomatal density are mainly associated with precipitation, while that of stomatal size is mainly associated with temperature, and the skewness and kurtosis of stomatal traits are less related to climatic and soil variables. Beyond mean climate variables, stomatal trait moments also vary with climatic seasonality and extreme conditions. Our findings extend the knowledge of stomatal trait–environment relationships to the ecosystem scale, with applications in predicting future water and carbon cycles.more » « less
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Summary A prevailing hypothesis posits that achieving higher maximum rates of leaf carbon gain and water loss is constrained by geometry and/or selection to limit the allocation of epidermal area to stomata (fS). Under this ‘stomatal‐area minimization hypothesis’, highergs,maxis associated with greater numbers of smaller stomata because this trait combination increasesgs,maxwith minimal increase infS, leading to relative conservation offSsemi‐independent ofgs,maxdue to coordination in stomatal size, density, and pore depth. An alternative hypothesis is that the evolution of highergs,maxcan be enabled by a greater epidermal area allocated to stomata, leading to positive covariation betweenfSandgs,max; we call this the ‘stomatal‐area adaptation hypothesis’. Under this hypothesis, the interspecific scaling betweengs,max, stomatal density, and stomatal size is a by‐product of selection on a moving optimalgs,max.We integrated biophysical and evolutionary quantitative genetic modeling with phylogenetic comparative analyses of a global data set of stomatal density and size from 2408 vascular forest species. The models present specific assumptions of both hypotheses and deduce predictions that can be evaluated with our empirical analyses of forest plants.There are three main results. First, neither the stomatal‐area minimization nor adaptation hypothesis is sufficient to be supported. Second, estimates of interspecific scaling from common regression methods cannot reliably distinguish between hypotheses when stomatal size is bounded. Third, we reconcile both hypotheses with the data by including an additional assumption that stomatal size is bounded by a wide range and under selection; we refer to this synthetic hypothesis as the ‘stomatal adaptation + bounded size’ hypothesis.This study advances our understanding of scaling between stomatal size and density by mathematically describing specific assumptions of competing hypotheses, demonstrating that existing hypotheses are inconsistent with observations, and reconciling these hypotheses with phylogenetic comparative analyses by postulating a synthetic model of selection ongs,max,fS, and stomatal size.more » « lessFree, publicly-accessible full text available December 1, 2026
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Lawson, Tracy (Ed.)Abstract Shifts in stomatal trait distributions across contrasting environments and their linkage with ecosystem productivity at large spatial scales have been unclear. Here, we measured the maximum stomatal conductance (g), stomatal area fraction (f), and stomatal space-use efficiency (e, the ratio of g to f) of 800 plant species ranging from tropical to cold-temperate forests, and determined their values for community-weighted mean, variance, skewness, and kurtosis. We found that the community-weighted means of g and f were higher in drier sites, and thus, that drought ‘avoidance’ by water availability-driven growth pulses was the dominant mode of adaptation for communities at sites with low water availability. Additionally, the variance of g and f was also higher at arid sites, indicating greater functional niche differentiation, whereas that for e was lower, indicating the convergence in efficiency. When all other stomatal trait distributions were held constant, increasing kurtosis or decreasing skewness of g would improve ecosystem productivity, whereas f showed the opposite patterns, suggesting that the distributions of inter-related traits can play contrasting roles in regulating ecosystem productivity. These findings demonstrate the climatic trends of stomatal trait distributions and their significance in the prediction of ecosystem productivity.more » « less
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