Globally accelerating trends in societal development and human environmental impacts since the mid-twentieth century are known as the Great Acceleration and have been discussed as a key indicator of the onset of the Anthropocene epoch . While reports on ecological responses (for example, changes in species range or local extinctions) to the Great Acceleration are multiplying , it is unknown whether such biotic responses are undergoing a similar acceleration over time. This knowledge gap stems from the limited availability of time series data on biodiversity changes across large temporal and geographical extents. Here we use a dataset of repeated plant surveys from 302 mountain summits across Europe, spanning 145 years of observation, to assess the temporal trajectory of mountain biodiversity changes as a globally coherent imprint of the Anthropocene. We find a continent-wide acceleration in the rate of increase in plant species richness, with five times as much species enrichment between 2007 and 2016 as fifty years ago, between 1957 and 1966. This acceleration is strikingly synchronized with accelerated global warming and is not linked to alternative global change drivers. The accelerating increases in species richness on mountain summits across this broad spatial extent demonstrate that acceleration in climate-induced biotic change is occurring even in remote places on Earth, with potentially far-ranging consequences not only for biodiversity, but also for ecosystem functioning and services.

Rapid climate warming in the tundra biome has been linked to increasing shrub dominance 1-4 . Shrub expansion can modify climate by altering surface albedo, energy and water balance, and permafrost 2,5-8 , yet the drivers of shrub growth remain poorly understood. Dendroecological data consisting of multi-decadal time series of annual shrub growth provide an underused resource to explore climate-growth relationships. Here, we analyse circumpolar data from 37 Arctic and alpine sites in 9 countries, including 25 species, and ∼42,000 annual growth records from 1,821 individuals. Our analyses demonstrate that the sensitivity of shrub growth to climate was: (1) heterogeneous, with European sites showing greater summer temperature sensitivity than North American sites, and (2) higher at sites with greater soil moisture and for taller shrubs (for example, alders and willows) growing at their northern or upper elevational range edges. Across latitude, climate sensitivity of growth was greatest at the boundary between the Low and High Arctic, where permafrost is thawing 4 and most of the global permafrost soil carbon pool is stored 9 . The observed variation in climate-shrub growth relationships should be incorporated into Earth system models to improve future projections of climate change impacts across the tundra biome.The Arctic is warming more rapidly than lower latitudes owing to climate amplification involving temperature, water vapour, albedo and sea ice feedbacks 5,7 . Tundra ecosystems are thus predicted to respond more rapidly to climate change than other terrestrial ecosystems 4 . The tundra biome spans Arctic and alpine regions that have similar plant species pools and mean climates, yet vary in topography, seasonality, land cover and glaciation history. Concurrent with the recent high-latitude warming trend 7 , repeat photography and vegetation surveys have shown widespread expansion of shrubs 1-3 , characterized by increased canopy cover, height and abundance. However, climate warming 7 and shrub increase 2,10 have not occurred at all sites. Models predict that warming of 2-10 • C (ref. 11) could convert as much as half of current tundra to 'shrubland' by the end of the twenty-first century 8 , but the uniformity of the frequently cited relationship between climate

The ongoing climate change is characterized by increased temperatures and altered precipitation patterns. In addition, there has been an increase in both the frequency and intensity of extreme climatic events such as drought. Episodes of drought induce a series of interconnected effects, all of which have the potential to alter the carbon balance of forest ecosystems profoundly at different scales of plant organization and ecosystem functioning. During recent years, considerable progress has been made in the understanding of how aboveground parts of trees respond to drought and how these responses affect carbon assimilation. In contrast, processes of belowground parts are relatively underrepresented in research on climate change. In this review, we describe current knowledge about responses of tree roots to drought. Tree roots are capable of responding to drought through a variety of strategies that enable them to avoid and tolerate stress. Responses include root biomass adjustments, anatomical alterations, and physiological acclimations. The molecular mechanisms underlying these responses are characterized to some extent, and involve stress signaling and the induction of numerous genes, leading to the activation of tolerance pathways. In addition, mycorrhizas seem to play important protective roles. The current knowledge compiled in this review supports the view that tree roots are well equipped to withstand drought situations and maintain morphological and physiological functions as long as possible. Further, the reviewed literature demonstrates the important role of tree roots in the functioning of forest ecosystems and highlights the need for more research in this emerging field.

Human societies depend on an Earth System that operates within a constrained range of nutrient 68 availability, yet the recent trajectory of terrestrial nitrogen (N) availability is uncertain. 69 Examining patterns of foliar N concentrations ([N]) and isotope ratios (δ 15 N) from more than 42,000 samples acquired over years, here we show that foliar [N] declined by 8% and foliar δ 15 N declined by 0.8 -1.9 ‰. Examining patterns across different climate spaces, foliar δ 15 N declined across the entire range of MAT and MAP tested. These results suggest declines in N supply relative to plant demand at the global scale. In all, there are now multiple lines of evidence of declining N availability in many unfertilized terrestrial ecosystems, including declines in δ 15 N of tree rings and leaves from herbarium samples over the past 75-150 years. 76These patterns are consistent with the proposed consequences of elevated atmospheric CO 2 and longer growing seasons. These declines will limit future terrestrial C uptake and increase nutritional stress for herbivores. 235 much. Preventing these declines in N availability further emphasizes the need to reduce 236 anthropogenic CO 2 emissions.Data and code availability. The datasets generated during and/or analysed during the current study are available in the Dryad repository [link to be generated upon acceptance]. All code used for statistical analyses and figure generation are available on Dryad (XXX).

The rapidly warming temperatures in high-latitude and alpine regions have the potential to alter the phenology of Arctic and alpine plants, affecting processes ranging from food webs to ecosystem trace gas fluxes. The International Tundra Experiment (ITEX) was initiated in 1990 to evaluate the effects of expected rapid changes in temperature on tundra plant phenology, growth and community changes using experimental warming. Here, we used the ITEX control data to test the phenological responses to background temperature variation across sites spanning latitudinal and moisture gradients. The dataset overall did not show an advance in phenology; instead, temperature variability during the years sampled and an absence of warming at some sites resulted in mixed responses. Phenological transitions of high Arctic plants clearly occurred at lower heat sum thresholds than those of low Arctic and alpine plants. However, sensitivity to temperature change was similar among plants from the different climate zones. Plants of different communities and growth forms differed for some phenological responses. Heat sums associated with flowering and greening appear to have increased over time. These results point to a complex suite of changes in plant communities and ecosystem function in high latitudes and elevations as the climate warms.

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Summary1. Using experimental atmospheric CO 2 enrichment, we tested for tree growth stimulation at the high-elevation treeline, where there is overwhelming evidence that low temperature inhibits growth despite an adequate carbon supply. We exposed Larix decidua (European larch) and Pinus mugo ssp. uncinata (mountain pine) to 9 years of free-air CO 2 enrichment (FACE) in an in situ experiment at treeline in the Swiss Alps (2180 m a.s.l.). 2. Accounting for pre-treatment vigour of individual trees, tree ring increments throughout the experimental period were larger in Larix growing under elevated CO 2 but not in Pinus. The magnitude of the CO 2 response in Larix ring width varied over time, with a significant stimulation occurring in treatment years 3-7 (marginal in year 6). 3. After 9 years of treatment, leaf canopy cover, stem basal area and total new shoot production were overall greater in Larix trees growing under elevated CO 2 , whereas Pinus showed no such cumulative growth response. The Larix ring width response in years 3-7 could have caused the cumulative CO 2 effect on tree size even if no further stimulation occurred, so it remains unclear if responsiveness was sustained over the longer term. 4. Larix ring width was stimulated more by elevated CO 2 in years with relatively high spring temperatures and an early snowmelt date, suggesting that temperatures were less limiting in these years and greater benefit was gained from extra carbon assimilated under elevated CO 2 . The magnitude of CO 2 stimulation was also larger after relatively high temperatures and high solar radiation in the preceding growing season, perhaps reflecting gains due to larger carbon reserves. 5. Synthesis. Contrasting above-ground growth responses of two treeline tree species to elevated CO 2 concentrations suggest that Larix will have a competitive advantage over less responsive species, such as co-occurring Pinus, under future CO 2 concentrations. Stimulation of Larix growth might be especially pronounced in a future warmer climate.

Climate warming may alter ecosystem nitrogen (N) cycling by accelerating N transformations in the soil, and changes may be especially pronounced in cold regions characterized by N-poor ecosystems. We investigated N dynamics across the plant-soil continuum during 6 years of experimental soil warming (2007-2012; +4 °C) at a Swiss high-elevation treeline site (Stillberg, Davos; 2180 m a.s.l.) featuring Larix decidua and Pinus uncinata. In the soil, we observed considerable increases in the NH4+ pool size in the first years of warming (by >50%), but this effect declined over time. In contrast, dissolved organic nitrogen (DON) concentrations in soil solutions from the organic layer increased under warming, especially in later years (maximum of +45% in 2012), suggesting enhanced DON leaching from the main rooting zone. Throughout the experimental period, foliar N concentrations showed species-specific but small warming effects, whereas δ N values showed a sustained increase in warmed plots that was consistent for all species analysed. The estimated total plant N pool size at the end of the study was greater (+17%) in warmed plots with Pinus but not in those containing Larix, with responses driven by trees. Irrespective of plot tree species identity, warming led to an enhanced N pool size of Vaccinium dwarf shrubs, no change in that of Empetrum hermaphroditum (dwarf shrub) and forbs, and a reduction in that of grasses, nonvascular plants, and fine roots. In combination, higher foliar δ N values and the transient response in soil inorganic N indicate a persistent increase in plant-available N and greater cumulative plant N uptake in warmer soils. Overall, greater N availability and increased DON concentrations suggest an opening of the N cycle with global warming, which might contribute to growth stimulation of some plant species while simultaneously leading to greater N losses from treeline ecosystems and possibly other cold biomes.