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1. Current changes in shrub abundance in alpine and arctic tundra ecosystems are primarily driven by climate change. However, while taller shrub communities are expanding, dwarf shrub communities show reductions under climate warming, and the mechanisms driving the latter (such as warming temperatures or accelerated spring snowmelt) may be complex. 2. To determine and disentangle the response of a widespread arctic-alpine prostrate dwarf shrub to both climate warming and changes in snowmelt time, we investigated phenology, clonal and sexual reproduction, leaf size, wood tissue carbon balance and leaf damage in 480 patches of Salix herbacea, along its elevational and snowmelt microhabitat range over 3 years in a space-for-time substitution. 3. Earlier snowmelt was associated with longer phenological development periods, an increased likelihood of herbivory and fungal damage, lower stem density, smaller leaves and lower end-of-season wood reserve carbohydrates. Furthermore, while early snowmelt was associated with an increased proportion of flowering stems, the proportion of fruiting stems was not, as fruit set decreased significantly with earlier snowmelt. Warmer temperatures at lower elevations were associated with lower stem numbers and larger leaves. 4. Synthesis. Our study indicates that phenology, fitness proxies and fungal/insect damage of the dwarf shrub S. herbacea are strongly influenced by snowmelt timing, and that earlier spring snowmelt reduced performance in S. herbacea. The likely mechanisms for many of the observed patterns are related to adverse temperature conditions in the early growing season. Reductions in clonal (stem number) and sexual reproduction (reduced fruit set) under earlier snowmelt, in addition to increasing damage probability, will likely lead to lower fitness and poorer performance, particularly in shrubs growing in early-exposure microhabitats. Further, we saw few concurrent benefits of higher temperatures for S. herbacea, particularly as warming was associated with lower clonal growth. As growing seasons become warmer and longer in arctic and alpine tundra ecosystems, early snowmelt is a critical mechanism reducing fitness and performance in a widespread dwarf shrub and may ultimately reduce dwarf shrub communities in tundra biomes.

Carbohydrate reserves play a vital role in plant survival during periods of negative carbon balance. Under a carbon-limited scenario, we expect a trade-offs between carbon allocation to growth, reserves, and defense. A resulting hypothesis is that carbon allocation to reserves exhibits a coordinated variation with functional traits associated with the ‘fast-slow’ plant economics spectrum. We tested the relationship between non-structural carbohydrates (NSC) of tree organs and functional traits using 61 angiosperm tree species from temperate and tropical forests with phylogenetic hierarchical Bayesian models. Our results provide evidence that NSC concentrations in stems and branches are decoupled from plant functional traits. while those in roots are weakly coupled with plant functional traits. In contrast, we found that variation between NSC concentrations in leaves and the fast-slow trait spectrum was coordinated, as species with higher leaf NSC had trait values associated with resource conservative species, such as lower SLA, leaf N, and leaf P. We also detected a small effect of leaf habit on the variation of NSC concentrations in branches and roots. Efforts to predict the response of ecosystems to global change will need to integrate a suite of plant traits, such as NSC concentrations in woody organs, that are independent of the ‘fast-slow’ plant economics spectrum and that capture how species respond to a broad range of global change drivers.

Climate change is altering spring snowmelt patterns in alpine and arctic ecosystems, and these changes may alter plant phenology, growth and reproduction. To predict how alpine plants respond to shifts in snowmelt timing, we need to understand trait plasticity, its effects on growth and reproduction, and the degree to which plants experience a home-site advantage. We tested how the common, long-lived dwarf shrub Salix herbacea responded to changing spring snowmelt time by reciprocally transplanting turfs of S. herbacea between early-exposure ridge and late-exposure snowbed microhabitats. After the transplant, we monitored phenological, morphological and fitness traits, as well as leaf damage, during two growing seasons. Salix herbacea leafed out earlier, but had a longer development time and produced smaller leaves on ridges relative to snowbeds. Longer phenological development times and smaller leaves were associated with reduced sexual reproduction on ridges. On snowbeds, larger leaves and intermediate development times were associated with increased clonal reproduction. Clonal and sexual reproduction showed no response to altered snowmelt time. We found no home-site advantage in terms of sexual and clonal reproduction. Leaf damage probability depended on snowmelt and thus exposure period, but had no short-term effect on fitness traits. We conclude that the studied populations of S. herbacea can respond to shifts in snowmelt by plastic changes in phenology and leaf size, while maintaining levels of clonal and sexual reproduction. The lack of a home-site advantage suggests that S. herbacea may not be adapted to different microhabitats. The studied populations are thus unlikely to react to climate change by rapid adaptation, but their responses will also not be constrained by small-scale local adaptation. In the short term, snowbed plants may persist due to high stem densities. However, in the long term, reduction in leaf size and flowering, a longer phenological development time and increased exposure to damage may decrease overall performance of S. herbacea under earlier snowmelt.

Alpine ecosystems are seriously threatened by climate change. One of the key mechanisms by which plants can adapt to changing environmental conditions is through evolutionary change. However, we still know little about the evolutionary potential in wild populations of long‐lived alpine plants. Here, we investigated heritabilities of phenological traits, leaf size, and performance traits in natural populations of the long‐lived alpine dwarf shrub Salix herbacea using relatedness estimates inferred from SSR (Simple Sequence Repeat) markers. Salix herbacea occurs in early‐ and late‐snowmelt microhabitats (ridges and snowbeds), and we assessed how performance consequences of phenological traits and leaf size differ between these microhabitats in order to infer potential for evolutionary responses. Salix herbacea showed low, but significant, heritabilities of leaf size, clonal and sexual reproduction, and moderate heritabilities of phenological traits. In both microhabitats, we found that larger leaves, longer intervals between snowmelt and leaf expansion, and longer GDD (growing‐degree days) until leaf expansion resulted in a stronger increase in the number of stems (clonal reproduction). In snowbeds, clonal reproduction increased with a shorter GDD until flowering, while the opposite was found on ridges. Furthermore, the proportion of flowering stems increased with GDD until flowering in both microhabitats. Our results suggest that the presence of significant heritable variation in morphology and phenology might help S. herbacea to adapt to changing environmental conditions. However, it remains to be seen if the rate of such an evolutionary response can keep pace with the rapid rate of climate change. We investigated heritabilities of phenological traits, leaf size, and performance traits in natural populations of the long‐lived alpine dwarf shrub Salix herbacea using relatedness estimates inferred from SSR markers. Our results suggest that the presence of significant heritable variation in morphology and phenology might help S. herbacea to adapt to changing environmental conditions.

Commentary p 264

Plants store large amounts of non-structural carbohydrates (NSC). While multiple functions of NSC have long been recognized, the interpretation of NSC seasonal dynamics is often based on the idea that stored NSC is a reservoir of carbon that fluctuates depending on the balance between supply via photosynthesis and demand for growth and respiration (the source–sink dynamics concept). Consequently, relatively high NSC concentrations in some plants have been interpreted to reflect excess supply relative to demand. An alternative view, however, is that NSC accumulation reflects the relatively high NSC levels required for plant survival; an important issue that remains highly controversial. Here, we assembled a new global database to examine broad patterns of seasonal NSC variation across organs (leaves, stems, and belowground), plant functional types (coniferous, drought-deciduous angiosperms, winter deciduous angiosperms, evergreen angiosperms, and herbaceous) and biomes (boreal, temperate, Mediterranean, and tropical). We compiled data from 121 studies, including seasonal measurements for 177 species under natural conditions. Our results showed that, on average, NSC account for ~10% of dry plant biomass and are highest in leaves and lowest in stems, whereas belowground organs show intermediate concentrations. Total NSC, starch, and soluble sugars (SS) varied seasonally, with a strong depletion of starch during the growing season and a general increase during winter months, particularly in boreal and temperate biomes. Across functional types, NSC concentrations were highest and most variable in herbaceous species and in conifer needles. Conifers showed the lowest stem and belowground NSC concentrations. Minimum NSC values were relatively high (46% of seasonal maximums on average for total NSC) and, in contrast to average values, were similar among biomes and functional types. Overall, although starch depletion was relatively common, seasonal depletion of total NSC or SS was rare. These results are consistent with a dual view of NSC function: whereas starch acts mostly as a reservoir for future use, soluble sugars perform immediate functions (e.g., osmoregulation) and are kept above some critical threshold. If confirmed, this dual function of NSC will have important implications for the way we understand and model plant carbon allocation and survival under stress.

Aim: Across all latitudes, high-elevation tree lines represent a drastic change in the dominant plant life-form, from upright trees to low-stature alpine plants. Although associated with low temperatures, the physiological mechanisms controlling this boundary are still not clear. The growth-limitation hypothesis assumes a direct low-temperature restriction of tissue formation at otherwise sufficient photoassimilation. In order to test this hypothesis, we present a global synthesis of previously published and new data on tree carbon supply status at high-elevation tree lines. Location: Global; 13 regions between 68° N and 45° S. Methods: Late-season concentrations of non-structural carbohydrates (NSC) in foliage and branch wood were measured at three elevations across the tree line ecotones, from upper tall forests (timber line) to the edges of aborescent tree growth (tree line). Year-round records of -10 cm soil temperatures were taken at the tree line. Results: Despite large differences in elevation and season length, the mean growing season temperature at the tree line was similar (approximately 6.6 °C) between all sites. NSC concentrations were not depleted at any of the elevation gradients between timber line and tree line, indicating no shortage of C at the uppermost tree stands. Tested across all sites, NSC concentrations in fact significantly increased with elevation. On average, NSC increased by 18% in leaves and 26% in branch wood from the lowest to the highest stands, primarily due to higher starch concentrations rather than free sugars. Hence, these responses do not reflect osmotic adjustments to lower temperatures at the end of the growing season. Main conclusions: This global data set contributes to a mechanistic understanding of tree line formation based on biological principles across climatic zones and tree genera. No evidence of C shortage was found at the high-elevation tree line in either seasonal and non-seasonal regions. The increasing trend of NSC concentrations with elevation is in line with the growth-limitation hypothesis.

Tree internal carbon (C) fluxes between compound and compartment pools are difficult to measure directly. Here we used a C mass balance approach to decipher these fluxes and provide a full description of tree C allocation dynamics. We collected independent measurements of tree C sinks, source and pools in Pinus halepensis in a semi‐arid forest, and converted all fluxes to g C per tree d⁻¹. Using this data set, a process flowchart was created to describe and quantify the tree C allocation on diurnal to annual time‐scales. The annual C source of 24.5 kg C per tree yr⁻¹was balanced by C sinks of 23.5 kg C per tree yr⁻¹, which partitioned into 70%, 17% and 13% between respiration, growth, and litter (plus export to soil), respectively. Large imbalances (up to 57 g C per tree d⁻¹) were observed as C excess during the wet season, and as C deficit during the dry season. Concurrent changes in C reserves (starch) were sufficient to buffer these transient C imbalances. The C pool dynamics calculated using the flowchart were in general agreement with the observed pool sizes, providing confidence regarding our estimations of the timing, magnitude, and direction of the internal C fluxes.

The water status of transpiring tree crowns depends on a hydraulic continuum from the soil matrix around roots to the sub-stomatal cavity of leaves, with a multitude of hydraulic resistances along this path. Although the stem xylem path may not be the most critical of these resistances, it had been suggested that a >50% interruption of that path by drought-stress-induced embolization (air filling) of conduits is critical for tree survival. Here we show that cutting the sapwood of mature, 35 m tall trees in half hardly affects crown water status and transpiration. Counter expectation, this first adult tree sapwood interception experiment revealed that shoot water potential in the canopy (assessed by using a 45 m canopy crane) either remained unaffected (spruce) or became less negative (beech), associated with small reductions in leaf diffusive conductance for water vapour. We conclude that the stem xylem of these trees has a large overcapacity and the tree hydraulics debate requires a critical re-visitation.