Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere–atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of droughtinduced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function

Because of the characteristically low temperatures and ambient CO 2 concentrations associated with greater altitudes, mountain forests may be particularly sensitive to global warming and increased atmospheric CO 2 . Moreover, the upper treeline is probably the most stressful location within these forests, possibly providing an early bellwether of forest response. Most treeline studies of the past century, as well as recently, have correlated temperatures with the altitudinal limits observed for treelines. In contrast, investigations on pre-establishment seedlings, the most vulnerable life stage of most tree species, are rare. There appears to be specific microclimatic factors dictated by wind and sky exposure that limit seedling survival, and also generate the distorted tree forms commonly observed at treeline. Seedling survival appears critical for creating the biological facilitation of microclimate at the community level which is necessary for the growth of seedlings to normal tree stature, forming new subalpine forest at a higher altitude.Abstract Es posible que-a causa de características que están asociadas con altitudes más altas: las bajas temperaturas y las concentraciones ambientales de dióxido de carbono-los bosques en las montañas están extra sensibles al calentamiento global y el aumento de dióxido de carbono en la atmósfera. El borde superior del bosque es probablemente el lugar con la más estrés y proviene uno de los primeros avisos de cómo reaccionará el bosque entero. En el pasado y hoy en día, la mayoría de los estudios del borde del bosque ha conectado la temperatura con los límites de la altitud. En contraste, investigaciones de árboles infantiles son raras, y la infancia de los árboles es el período de vida más vulnerable. Aparece que hay factores micro-climáticos dictados por la exposición del viento y cielo que limitan la sobrevivencia de los árboles infantiles, y que generan árboles deformados observados al borde del bosque. Es más, la sobrevivencia de árboles infantiles es crítica para crear la facilitación biológica del micro-clima en una comunidad arbolada. Esta facilitación es necesaria para el crecimiento de árboles infantiles a árboles maduros, los que forman un nuevo bosque subalpino en una altitud más alta.

Summary1. Ecohydrological niches are important for understanding plant community responses to climate shifts, particularly in dry lands. According to the two-layer hypothesis, selective use of deep-soil water increases growth or persistence of woody species during warm and dry summer periods and thereby contributes to their coexistence with shallow-rooted herbs in dry ecosystems. The resourcepool hypothesis further suggests that shallow-soil water benefits growth of all plants while deep-soil water primarily enhances physiological maintenance and survival of woody species. Few studies have directly tested these by manipulating deep-soil water availability and observing the long-term outcomes. 2. We predicted that factors promoting infiltration and storage of water in deep soils, specifically greater winter precipitation and soil depth, would enhance Artemisia tridentata (big sagebrush) in cold, winter-wet/summer-dry desert. Sagebrush responses to 20 years of winter irrigation were compared to summer-or no irrigation, on plots having relatively deep or shallow soils (2 m vs. 1 m depths). 3. Winter irrigation increased sagebrush cover, and crown and canopy volumes, but not density (individuals/plot) compared to summer or no irrigation, on deep-soil plots. On shallow-soil plots, winter irrigation surprisingly decreased shrub cover and size, and summer irrigation had no effect. Furthermore, multiple regression suggested that the variations in growth were related (i) firstly to water in shallow soils (0-0.2 m) and secondly to deeper soils (> 1 m deep) and (ii) more by springtime than by midsummer soil water. Water-use efficiency increased considerably on shallow soils without irrigation and was lowest with winter irrigation. 4. Synthesis. Sagebrush was more responsive to the seasonal timing of precipitation than to total annual precipitation. Factors that enhanced deep-water storage (deeper soils plus more winter precipitation) led to increases in Artemisia tridentata that were consistent with the two-layer hypothesis, and the contribution of shallow water to growth on these plots was consistent with the resource-pool hypothesis. However, shallow-soil water also had negative effects on sagebrush, suggesting an ecohydrological trade-off not considered in these or related theories. The interaction between precipitation timing and soil depth indicates that increased winter precipitation could lead to a mosaic of increases and decreases in A. tridentata across landscapes having variable soil depth.

No abstract

The spatial patterning of alpine plant communities is strongly influenced by the variation in physical factors such as temperature and moisture, which are strongly affected by snow depth and snowmelt patterns. Earlier snowmelt timing and greater soil-moisture limitations may favor wide-ranging species adapted to a broader set of ecohydrological conditions than alpine-restricted species. We asked how plant community composition, phenology, plant water relations, and photosynthetic gas exchange of alpine-restricted and wide-ranging species differ in their responses to a ca. 40-day snowmelt gradient in the Colorado Rocky Mountains (Lewisia pygmaea, Sibbaldia procumbens, and Hymenoxys grandiflora were alpine-restricted and Artemisia scopulorum, Carex rupestris, and Geum rossii were wide-ranging species). As hypothesized, species richness and foliar cover increased with earlier snowmelt, due to a greater abundance of wide-ranging species present in earlier melting plots. Flowering initiation occurred earlier with earlier snowmelt for 12 out of 19 species analyzed, while flowering duration was shortened with later snowmelt for six species (all but one were wide-ranging species). We observed >50% declines in net photosynthesis from July to September as soil moisture and plant water potentials declined. Early-season stomatal conductance was higher in wide-ranging species, indicating a more competitive strategy for water acquisition when soil moisture is high. Even so, there were no associated differences in photosynthesis or transpiration, suggesting no strong differences between these groups in physiology. Our findings reveal that plant species with different ranges (alpine-restricted vs. wide-ranging) could have differential phenological and physiological responses to snowmelt timing and associated soil moisture dry-down, and that alpine-restricted species’ performance is more sensitive to snowmelt. As a result, alpine-restricted species may serve as better indicator species than their wide-ranging heterospecifics. Overall, alpine community composition and peak % cover are strongly structured by spatio-temporal patterns in snowmelt timing. Thus, near-term, community-wide changes (or variation) in phenology and physiology in response to shifts in snowmelt timing or rates of soil dry down are likely to be contingent on the legacy of past climate on community structure.