Recent research using repeat photography, long-term ecological monitoring and dendrochronology has documented shrub expansion in arctic, high-latitude and alpine tundra 1 1748-9326/11/045509+15$33.00 c 2011 IOP Publishing Ltd Printed in the UK Environ. Res. Lett. 6 (2011) 045509 I H Myers-Smith et al ecosystems.Here, we (1) synthesize these findings, (2) present a conceptual framework that identifies mechanisms and constraints on shrub increase, (3) explore causes, feedbacks and implications of the increased shrub cover in tundra ecosystems, and (4) address potential lines of investigation for future research. Satellite observations from around the circumpolar Arctic, showing increased productivity, measured as changes in 'greenness', have coincided with a general rise in high-latitude air temperatures and have been partly attributed to increases in shrub cover. Studies indicate that warming temperatures, changes in snow cover, altered disturbance regimes as a result of permafrost thaw, tundra fires, and anthropogenic activities or changes in herbivory intensity are all contributing to observed changes in shrub abundance. A large-scale increase in shrub cover will change the structure of tundra ecosystems and alter energy fluxes, regional climate, soil-atmosphere exchange of water, carbon and nutrients, and ecological interactions between species. In order to project future rates of shrub expansion and understand the feedbacks to ecosystem and climate processes, future research should investigate the species or trait-specific responses of shrubs to climate change including: (1) the temperature sensitivity of shrub growth, (2) factors controlling the recruitment of new individuals, and (3) the relative influence of the positive and negative feedbacks involved in shrub expansion.

Here we present a novel, empirical approach to assess the relative sensitivity of ecosystems to climate variability, one property of resilience that builds on theoretical modelling work recognizing that systems closer to critical thresholds respond more sensitively to external perturbations 2 . We develop a new metric, the vegetation sensitivity index (VSI) which identifies areas sensitive to climate variability over the past 14 years. The metric uses time series data of MODIS-derived enhanced vegetation index (EVI) 3and three climatic variables that drive vegetation productivity 4 (air temperature, water availability and cloud cover). Underlying the analysis is an autoregressive modelling approach used to identify climate drivers of vegetation productivity on monthly timescales, in addition to regions with memory effects and reduced response rates to external forcing 5 . We find ecologically sensitive regions with amplified responses to climate variability in the arctic tundra, parts of the boreal forest belt, the tropical rainforest, alpine regions worldwide, steppe and prairie regions of central Asia and North and South America, the Caatinga deciduous forest in eastern South America, and eastern areas of Australia. Our study provides a quantitative methodology for assessing the relative response rate of ecosystems-be they natural or with a strong anthropogenic signature-to environmental variability, which is the first step toward addressing why some regions appear to be more sensitive than others and what impact this has upon the resilience of ecosystem service provision and human well-being.The rate and scale of projected climate changes in the 21st century are likely to have profound impacts on the functioning of Earth's ecosystems 6 . Much current understanding of how biodiversity will respond to climate change is based on responses to changes in mean Publisher: NPG; Journal: Nature: Nature; Article Type: Biology letter DOI: 10.1038/nature16986Page 2 of 25 climate state 7 . However, climate variability, and the related increases in extreme events in a warmer world 8 , has a strong influence on both the structuring and functioning of ecosystems [9][10][11] . Given the importance of identifying ecologically sensitive areas for ecosystem service provision and poverty alleviation 1 , a key knowledge gap exists in how to identify and then prioritize those regions that are most sensitive to climatic variability.Ecosystem response to variability in external forcing is a key component of resilience.Theory indicates that systems with lower resilience (that is, those with a high probability of crossing a threshold to an alternative state 12 ) experience amplified responses to disturbance and are more sensitive to environmental perturbations 2 . In addition, slower responses (identified through increased autocorrelation) may be evidence of reduced recovery rates in systems approaching critical transitions 13 . Therefore, identification of areas with high ecological sensitivity or reduced recovery rates is an importan...

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

2020). Complexity revealed in the greening of the Arctic. Nature Climate Change, 10 pp. 106-117.For guidance on citations see FAQs.

Growth in arctic vegetation is generally expected to increase under a warming climate, particularly among deciduous shrubs. We analyzed annual ring growth for an abundant and nearly circumpolar erect willow (Salix lanata L.) from the coastal zone of the northwest Russian Arctic (Nenets Autonomous Okrug). The resulting chronology is strongly related to summer temperature for the period 1942-2005. Remarkably high correlations occur at long distances (41600 km) across the tundra and taiga zones of West Siberia and Eastern Europe. We also found a clear relationship with photosynthetic activity for upland vegetation at a regional scale for the period 1981-2005, confirming a parallel 'greening' trend reported for similarly warming North American portions of the tundra biome. The standardized growth curve suggests a significant increase in shrub willow growth over the last six decades. These findings are in line with field and remote sensing studies that have assigned a strong shrub component to the reported greening signal since the early 1980s. Furthermore, the growth trend agrees with qualitative observations by nomadic Nenets reindeer herders of recent increases in willow size in the region. The quality of the chronology as a climate proxy is exceptional. Given its wide geographic distribution and the ready preservation of wood in permafrost, S. lanata L. has great potential for extended temperature reconstructions in remote areas across the Arctic.

Over the past decade, the Arctic has warmed by 0.75°C, far outpacing the global average, while Antarctic temperatures have remained comparatively stable. As Earth approaches 2°C warming, the Arctic and Antarctic may reach 4°C and 2°C mean annual warming, and 7°C and 3°C winter warming, respectively. Expected consequences of increased Arctic warming include ongoing loss of land and sea ice, threats to wildlife and traditional human livelihoods, increased methane emissions, and extreme weather at lower latitudes. With low biodiversity, Antarctic ecosystems may be vulnerable to state shifts and species invasions. Land ice loss in both regions will contribute substantially to global sea level rise, with up to 3 m rise possible if certain thresholds are crossed. Mitigation efforts can slow or reduce warming, but without them northern high latitude warming may accelerate in the next two to four decades. International cooperation will be crucial to foreseeing and adapting to expected changes.

Arctic warming can influence tundra ecosystem function with consequences for climate feedbacks, wildlife and human communities. Yet ecological change across the Arctic tundra biome remains poorly quantified due to field measurement limitations and reliance on coarse-resolution satellite data. Here, we assess decadal changes in Arctic tundra greenness using time series from the 30 m resolution Landsat satellites. From 1985 to 2016 tundra greenness increased (greening) at ~37.3% of sampling sites and decreased (browning) at ~4.7% of sampling sites. Greening occurred most often at warm sampling sites with increased summer air temperature, soil temperature, and soil moisture, while browning occurred most often at cold sampling sites that cooled and dried. Tundra greenness was positively correlated with graminoid, shrub, and ecosystem productivity measured at field sites. Our results support the hypothesis that summer warming stimulated plant productivity across much, but not all, of the Arctic tundra biome during recent decades.

Summary1. Priority question exercises are becoming an increasingly common tool to frame future agendas in conservation and ecological science. They are an effective way to identify research foci that advance the field and that also have high policy and conservation relevance. 2. To date, there has been no coherent synthesis of key questions and priority research areas for palaeoecology, which combines biological, geochemical and molecular techniques in order to reconstruct past ecological and environmental systems on time-scales from decades to millions of years. 3. We adapted a well-established methodology to identify 50 priority research questions in palaeoecology. Using a set of criteria designed to identify realistic and achievable research goals, we selected questions from a pool submitted by the international palaeoecology research community and relevant policy practitioners. 4. The integration of online participation, both before and during the workshop, increased international engagement in question selection. 5. The questions selected are structured around six themes: human-environment interactions in the Anthropocene; biodiversity, conservation and novel ecosystems; biodiversity over long time-scales; ecosystem processes and biogeochemical cycling; comparing, combining and synthesizing information from multiple records; and new developments in palaeoecology. 6. Future opportunities in palaeoecology are related to improved incorporation of uncertainty into reconstructions, an enhanced understanding of ecological and evolutionary dynamics and processes and the continued application of long-term data for better-informed landscape management. 256-26750 priority research questions in palaeoecology 257 7. Synthesis. Palaeoecology is a vibrant and thriving discipline, and these 50 priority questions highlight its potential for addressing both pure (e.g. ecological and evolutionary, methodological) and applied (e.g. environmental and conservation) issues related to ecological science and global change.