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.

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Passive open-top devices have been proposed as a method to experimentally increase temperature in high-latitude ecosystems. There is, however, little documentation on the efficacy of these devices. This paper examines the performance of four open-top chambers for altering temperature at six sites in the Arctic and Antarctica. Most of the heating effect was due to daytime warming above ambient; occasional night-time cooling below ambient, especially of air temperatures, depressed mean daily temperature. The mean daily temperatures at four arctic sites were generally increased by 1.2-1.8°C; but occasionally, temperature depressions also occurred. Under optimal conditions at the antarctic site (dry soils, no vegetation, high radiation) mean daily soil temperatures were increased by ⍣2.2°C (-10 cm) to ⍣5.2°C (0 cm). Protection from wind may play a more important role than temperature per se in providing a favourable environment for plant growth within opentop devices. Wind speed had a generally negative impact on mean daily temperature. Daily global radiation was both positively and negatively related to chamber temperature response. The effect of chambers on snow accumulation was variable with the Alexandra Fjord site showing an increased accumulation in chambers but no difference in the date of snowmelt, while at Latnjajaure in a deep snowfall site, snowmelt occurred 1-2 weeks earlier in chambers, potentially increasing the growing season. Selection of a passive temperature-enhancing system requires balancing the temperature enhancement desired against potential unwanted ecological effects such as chamber overheating and altered light, moisture, and wind. In general, the more closed the temperature-enhancing system, the higher is the temperature enhancement, but the larger are the unwanted ecological effects. Open-top chambers alter temperature significantly and minimize most unwanted ecological effects; as a consequence, these chambers are a useful tool for studying the response of high-latitude ecosystems to warming.

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