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.

Summary• In a comparative study of 42 rainforest tree species we examined relationships amongst wood traits, diameter growth and survival of large trees in the field, and shade tolerance and adult stature of the species.• The species show two orthogonal axes of trait variation: a primary axis related to the vessel size-number trade-off (reflecting investment in hydraulic conductance vs hydraulic safety) and a secondary axis related to investment in parenchyma vs fibres (storage vs strength). Across species, growth rate was positively related to vessel diameter and potential specific hydraulic conductance (K p ), and negatively related to wood density. Survival rate was only positively related to wood density.• Light-demanding species were characterized by low wood and vessel density and wide vessels. Tall species were characterized by wide vessels with low density and large K p . Hydraulic traits were more closely associated with adult stature than with light demand, possibly because tall canopy species experience more drought stress and face a higher cavitation risk.• Vessel traits affect growth and wood density affects growth and survival of large trees in the field. Vessel traits and wood density are therefore important components of the performance and life history strategies of tropical tree species.

Summary• The variability of branch-level hydraulic properties was assessed across 12 Scots pine populations covering a wide range of environmental conditions, including some of the southernmost populations of the species. The aims were to relate this variability to differences in climate, and to study the potential tradeoffs between traits.• Traits measured included wood density, radial growth, xylem anatomy, sapwoodand leaf-specific hydraulic conductivity (K S and K L ), vulnerability to embolism, leaf-to-sapwood area ratio (A L : A S ), needle carbon isotope discrimination (Δ 13 C) and nitrogen content, and specific leaf area.• Between-population variability was high for most of the hydraulic traits studied, but it was directly associated with climate dryness (defined as a combination of atmospheric moisture demand and availability) only for A L : A S , K L and Δ 13 C. Shoot radial growth and A L : A S declined with stand development, which is consistent with a strategy to avoid exceedingly low water potentials as tree size increases. In addition, we did not find evidence at the intraspecific level of some associations between hydraulic traits that have been commonly reported across species.• The adjustment of Scots pine's hydraulic system to local climatic conditions occurred primarily through modifications of A L : A S and direct stomatal control, whereas intraspecific variation in vulnerability to embolism and leaf physiology appears to be limited.

SummaryVariability in xylem anatomy is of interest to plant scientists because of the role water transport plays in plant performance and survival. Insights into plant adjustments to changing environmental conditions have mainly been obtained through structural and functional comparative studies between taxa or within taxa on contrasting sites or along environmental gradients. Yet, a gap exists regarding the study of hydraulic adjustments in response to environmental changes over the lifetimes of plants. In trees, dated tree-ring series are often exploited to reconstruct dynamics in ecological conditions, and recent work in which wood-anatomical variables have been used in dendrochronology has produced promising results. Environmental signals identified in water-conducting cells carry novel information reflecting changes in regional conditions and are mostly related to short, subannual intervals. Although the idea of investigating environmental signals through wood anatomical time series goes back to the 1960s, it is only recently that lowcost computerized image-analysis systems have enabled increased scientific output in this field. We believe that the study of tree-ring anatomy is emerging as a promising approach in tree biology and climate change research, particularly if complemented by physiological and ecological studies. This contribution presents the rationale, the potential, and the methodological challenges of this innovative approach.

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.