Plant functional trait change across a warming tundra biomeThe tundra is warming more rapidly than any other biome on Earth, and the potential ramifications are far-reaching because of global feedback effects between vegetation and climate. A better understanding of how environmental factors shape plant structure and function is crucial for predicting the consequences of environmental change for ecosystem functioning. Here we explore the biome-wide relationships between temperature, moisture and seven key plant functional traits both across space and over three decades of warming at 117 tundra locations. Spatial temperature-trait relationships were generally strong but soil moisture had a marked influence on the strength and direction of these relationships, highlighting the potentially important influence of changes in water availability on future trait shifts in tundra plant communities. Community height increased with warming across all sites over the past three decades, but other traits lagged far behind predicted rates of change. Our findings highlight the challenge of using space-for-time substitution to predict the functional consequences of future warming and suggest that functions that are tied closely to plant height will experience the most rapid change. They also reveal the strength with which environmental factors shape biotic communities at the coldest extremes of the planet and will help to improve projections of functional changes in tundra ecosystems with climate warming. Environment-trait relationships across the tundra biomeWe found strong spatial associations between temperature and community height, SLA and LDMC (Fig. 2a, Extended Data Fig. 2 and Supplementary Table 3) across the 117 survey sites. Both height and SLA increased with summer temperature, but the temperaturetrait relationship for SLA was much stronger at wetter than at drier sites. LDMC was negatively related to temperature, and

Part of the missing sink in the global CO 2 budget has been attributed to the positive effects of CO 2 fertilization and N deposition on carbon sequestration in Northern Hemisphere terrestrial ecosystems. The genus Sphagnum is one of the most important groups of plant species sequestrating carbon in temperate and northern bog ecosystems, because of the low decomposability of the dead material it produces. The effects of raised CO 2 and increased atmospheric N deposition on growth of Sphagnum and other plants were studied in bogs at four sites across Western Europe. Contrary to expectations, elevated CO 2 did not signi®cantly affect Sphagnum biomass growth. Increased N deposition reduced Sphagnum mass growth, because it increased the cover of vascular plants and the tall moss Polytrichum strictum. Such changes in plant species composition may decrease carbon sequestration in Sphagnum-dominated bog ecosystems.

Climate change is expected to cause extensive vegetation changes in the Arctic: deciduous shrubs are already expanding, in response to climate warming. The results from transect studies suggest that increasing shrub cover will impact significantly on the surface energy balance. However, little is known about the direct effects of shrub cover on permafrost thaw during summer. We experimentally quantified the influence of Betula nana cover on permafrost thaw in a moist tundra site in northeast Siberia with continuous permafrost. We measured the thaw depth of the soil, also called the active layer thickness (ALT), ground heat flux and net radiation in 10 m diameter plots with natural B. nana cover (control plots) and in plots in which B. nana was removed (removal plots). Removal of B. nana increased ALT by 9% on average late in the growing season, compared with control plots. Differences in ALT correlated well with differences in ground heat flux between the control plots and B. nana removal plots. In the undisturbed control plots, we found an inverse correlation between B. nana cover and late growing season ALT. These results suggest that the expected expansion of deciduous shrubs in the Arctic region, triggered by climate warming, may reduce summer permafrost thaw. Increased shrub growth may thus partially offset further permafrost degradation by future temperature increases. Permafrost models need to include a dynamic vegetation component to accurately predict future permafrost thaw.

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Abstract. Deciduous shrubs are expected to rapidly expand in the Arctic during the coming decades due to climate warming. A transition towards more shrub-dominated tundra may have large implications for the regional surface energy balance, permafrost stability and carbon storage capacity, with consequences for the global climate system. However, little information is available on the natural long-term shrub growth response to climatic variability. Our aim was to determine the climate factor and time period that are most important to annual shrub growth in our research site in NESiberia. Therefore, we determined annual radial growth rates in Salix pulchra and Betula nana shrubs by measuring ring widths. We constructed shrub ring width chronologies and compared growth rates to regional climate and remotely sensed greenness data. Early summer temperature was the most important factor influencing ring width of S. pulchra (Pearson's r = 0.73, p < 0.001) and B. nana (Pearson's r = 0.46, p < 0.001). No effect of winter precipitation on shrub growth was observed. In contrast, summer precipitation of the previous year correlated positively with B. nana ring width (Pearson's r = 0.42, p < 0.01), suggesting that wet summers facilitate shrub growth in the following growing season. S. pulchra ring width correlated positively with peak summer NDVI, despite the small coverage of S. pulchra shrubs (<5 % surface cover) in our research area. We provide the first climate-growth study on shrubs for Northeast Siberia, the largest tundra region in the world. We show thatCorrespondence to: D. Blok (daan.blok@wur.nl) two deciduous shrub species with markedly different growth forms have a similar growth response to changes in climate. The obtained shrub growth response to climate variability in the past increases our understanding of the mechanisms underlying current shrub expansion, which is required to predict future climate-driven tundra vegetation shifts.