The complex multi-sectoral impacts of drought: Evidence from a mountainous basin in the Central Spanish Pyrenees.

We analyzed the impacts of drought severity on a variety of sectors in a topographically complex basin (the upper Aragón basin 2181 km2) in the Central Spanish Pyrenees. Using diverse data sources including meteorological and hydrological observations, remote sensing and tree rings, we analyze the possible hydrological implications of drought occurrence and severity on water availability in various sectors, including downstream impacts on irrigation water supply for crop production. Results suggest varying responses in forest activity, secondary growth, plant phenology, and crop yield to drought impacts. Specifically, meteorological droughts have distinct impacts downstream, mainly due to water partitioning between streamflow and irrigation channels that transport water to crop producing areas. This implies that drought severity can extend beyond the physical boundaries of the basin, with impacts on crop productivity. This complex response to drought impacts makes it difficult to develop objective basin-scale operational definitions for monitoring drought severity. Moreover, given the high spatial variability in responses to drought across sectors, it is difficult to establish reliable drought thresholds from indices that are relevant across all socio-economic sectors. The anthropogenic impacts (e.g. water regulation projects, ecosystem services, land cover and land use changes) pose further challenges to assessing the response of different systems to drought severity. This study stresses the need to consider the seasonality of drought impacts and appropriate drought time scales to adequately assess and understand their complexity.

Mushroom productivity trends in relation to tree growth and climate across different European forest biomes.

Although it is logical to think that mycorrhizal mushroom production should be somehow related to the growth of the trees from which the fungi obtain carbohydrates, little is known about how mushroom yield patterns are related to tree performance. In this study, we delved into the understanding of the relationships between aboveground fungal productivity, tree radial growth patterns and climatic conditions across three latitudinally different bioclimatic regions encompassing Mediterranean, temperate and boreal forest ecosystems in Europe. For this purpose, we used a large assemblage of long-term data of weekly or biweekly mushroom yield monitoring in Spain, Switzerland and Finland. We analysed the relationships between annual mushroom yield (considering both biomass and number of sporocarps per unit area), tree ring features (tree ring, earlywood and latewood widths), and meteorological conditions (i.e. precipitation and temperature of summer and autumn) from different study sites and forest ecosystems, using both standard and partial correlations. Moreover, we fitted predictive models to estimate mushroom yield from mycorrhizal and saprotrophic fungal guilds based on climatic and dendrochronological variables. Significant synchronies between mushroom yield and climatic and dendrochronological variables were mostly found in drier Mediterranean sites, while few or no significant correlations were found in the boreal and temperate regions. We observed positive correlations between latewood growth and mycorrhizal mushroom biomass only in some Mediterranean sites, this relationship being mainly mediated by summer and autumn precipitation. Under more water-limited conditions, both the seasonal wood production and the mushroom yield are more sensitive to precipitation events, resulting in higher synchrony between both variables. This comparative study across diverse European forest biomes and types provides new insights into the relationship between mushroom productivity, tree growth and weather conditions.

Coexisting oak species, including rear-edge populations, buffer climate stress through xylem adjustments.

The ability of trees to cope with climate change is a pivotal feature of forest ecosystems, especially for rear-edge populations facing warm and dry conditions. To evaluate current and future forests threats, a multi-proxy focus on the growth, anatomical and physiological responses to climate change is needed. We examined the long-term xylem adjustments to climate variability of the temperate Quercus robur L. at its rear edge and the sub-Mediterranean Quercus pyrenaica Willd. Both species coexist at a mesic (ME, humid and warmer) and a xeric (XE, dry and cooler) site in northern Spain, the latter experiencing increasing temperatures in recent decades. We compared xylem traits at each site and assessed their trends, relationships and responses to climate (1960-2008). Traits included basal area increment, earlywood vessel hydraulic diameter, density and theoretical-specific hydraulic conductivity together with latewood oxygen (δ18O) stable isotopes and δ13C-derived water-use efficiency (iWUE). Quercus robur showed the highest growth at ME, likely through enhanced cambial activity. Quercus pyrenaica had higher iWUE at XE compared with ME, but limited plasticity of anatomical xylem traits was found for the two oak species. Similar physiological performance was found for both species. The iWUE augmented in recent years especially at XE, likely explained by stomatal closure given the increasing δ18O signal in response to drier and sunnier growing seasons. Overall, traits were more correlated at XE than at ME. The iWUE improvements were linked to higher growth up to a threshold (~85 µmol mol-1) after which reduced growth was found at XE. Our results are consistent with Q. pyrenaica and Q. robur coexisting at the central and dry edge of the climatic species distribution, respectively, showing similar responses to buffer warmer conditions. In fact, the observed adjustments found for Q. robur point towards growth stability of similar rear-edge oak populations under warmer climate conditions.

Winter drought impairs xylem phenology, anatomy and growth in Mediterranean Scots pine forests.

Continental Mediterranean forests face drought but also cold spells and both climate extremes can impair the resilience capacity of these forests. Climate warming could amplify the negative effects of cold spells by inducing premature dehardening. Here we capitalize on a winter drought-induced dieback triggered by a cold spell which occurred in December 2001 affecting Scots pine forests in eastern Spain. We assessed post-dieback recovery by quantifying and comparing radial growth and xylem anatomy of non-declining (ND, crown cover >50%) and declining (D, crown cover ≤50%) trees in two sites (VP, Villarroya de los Pinares; TO, Torrijas). We also characterized xylogenesis in both sites and aboveground productivity in site VP. Dieback caused legacy effects since needle loss, a 60% reduction in litter fall and radial-growth decline characterized D-trees 3 years after dieback symptoms started appearing in spring 2002. D-trees formed collapsed tracheids in the 2002-ring, particularly in the most affected VP site where xylogenesis differences between ND and D trees were most noticeable. The lower growth rates of D-trees were caused by a shorter duration of their major xylogenesis phases. In site VP the radial-enlargement and wall-thickening of tracheids were significantly reduced in D-trees as compared to ND-trees because these xylogenesis phases tended to start earlier and end later in ND-trees. Gompertz models fitted to tracheid production predicted that maximum growth rates occurred 11-12 days earlier in ND than in D-trees. The formation of radially-enlarging tracheids was enhanced by longer days in both study sites and also by wetter conditions in the driest TO site, but xylogenesis sensitivity to climate was reduced in D-trees. Winter-drought dieback impairs xylem anatomy and phenology, aboveground productivity, xylogenesis and growth in Mediterranean Scots pine populations. Affected stands show a costly post-dieback recovery challenging their resilience ability.

Topography and age mediate the growth responses of Smith fir to climate warming in the southeastern Tibetan Plateau.

The Tibetan Plateau holds some of the world's highest undisturbed natural treelines and timberlines. Such extreme environments constitute potentially valuable monitoring sites of the effects of climate warming on high-elevation forests. Here, we analyze a network of 21 Smith fir forests situated in the Sygera Mountains, southeastern Tibetan Plateau, using tree-ring width (TRW) and basal area increment (BAI) chronologies. Sampled sites encompassed a wide elevation gradient, from 3600 to 4400 m, including some treeline sites and diverse aspects and tree ages. In comparison with TRW series, BAI series better capture the long-term warming signal. Previous November and current April and summer temperatures are the dominant climatic factors controlling Smith fir radial growth. The mean inter-series correlations of TRW increased upwards, but the forest limit presented the highest potential to reconstruct past temperature variability. Moreover, the growth responses of young trees were less stable than those of trees older than 100 years. Climate warming is accelerating radial growth of Smith fir forest subjected to mesic conditions. Collectively, these findings confirm that the effects of site elevation and tree age should be considered when quantifying climate-growth relationships. The type of tree-ring data (BAI vs. TRW) is also relevant since BAI indices seem to be a better climatic proxy of low-frequency temperature signals than TRW indices. Therefore, site (e.g., elevation) and tree (e.g., age) features should be considered to properly evaluate the effects of climate warming on growth of high-elevation forests.

FOREST ECOLOGY. Pervasive drought legacies in forest ecosystems and their implications for carbon cycle models.

The impacts of climate extremes on terrestrial ecosystems are poorly understood but important for predicting carbon cycle feedbacks to climate change. Coupled climate-carbon cycle models typically assume that vegetation recovery from extreme drought is immediate and complete, which conflicts with the understanding of basic plant physiology. We examined the recovery of stem growth in trees after severe drought at 1338 forest sites across the globe, comprising 49,339 site-years, and compared the results with simulated recovery in climate-vegetation models. We found pervasive and substantial "legacy effects" of reduced growth and incomplete recovery for 1 to 4 years after severe drought. Legacy effects were most prevalent in dry ecosystems, among Pinaceae, and among species with low hydraulic safety margins. In contrast, limited or no legacy effects after drought were simulated by current climate-vegetation models. Our results highlight hysteresis in ecosystem-level carbon cycling and delayed recovery from climate extremes.

Contrasting seasonal overlaps between primary and secondary growth are linked to wood anatomy in Mediterranean sub-shrubs.

Whole-plant approaches allow quantification of the temporal overlap between primary and secondary growth. If the amount of time available to grow is short, there may be a high temporal overlap between shoot growth and wood formation. We hypothesise that such overlap depends on the duration of the growing season and relates to wood anatomy. We evaluated wood anatomy, shoot longitudinal and radial growth rates, fine root production and the concentrations of non-structural carbohydrates (NSC) in the wood of six sub-shrub species growing in sites with contrasting climatic conditions (Lepidium subulatum, Linum suffruticosum, Salvia lavandulifolia, Satureja montana, Ononis fruticosa, Echinospartum horridum). Sub-shrub species living in sites with a short growing season displayed a high overlap between aboveground primary and secondary growth and formed wide vessels, whereas species from the warmest and driest sites presented the reverse characteristics. The highest overlap was linked to a rapid shoot extension and thickening through the enhanced hydraulic conductivity provided by wide vessels. The reductions in NSC concentrations when growth peaked were low or moderate, indicating that sub-shrubs accumulate NSC in excess, as do trees. The temporal overlap among primary and secondary growth in woody plants may be connected to the duration and rates of shoot and wood growth, which in turn depend on the vessel lumen area.