Summary1. Some disturbances can drive ecological systems to abrupt shifts between alternative stages (tipping points) when critical transitions occur. Drought-induced tree death can be considered as a nonlinear shift in tree vigour and growth. However, at what point do trees become predisposed to drought-related dieback and which factors determine this (tipping) point? We investigated these questions by characterizing the responses of three tree species, silver fir (Abies alba), Scots pine (Pinus sylvestris) and Aleppo pine (Pinus halepensis), to a severe drought event. 2. We compared basal area increment (BAI) trends and responses to climate and drought in declining (very defoliated and dying) vs. non-declining (slightly or not defoliated) trees by using generalized additive mixed models. Defoliation, BAI and sapwood production were related to functional proxies of tree vigour measured at the onset and end of the drought (non-structural carbohydrate concentrations, needle N content and C isotopic discrimination, presence of wood-inhabiting fungi). We evaluated whether early warning signals (increases in synchronicity among trees or in autocorrelation and standard deviation) could be extracted from the BAI series prior to tree death. 3. Declining silver fir and Scots pine trees showed less growth than non-declining trees one to three decades, respectively, before the drought event, whereas Aleppo pines showed growth decline irrespective of tree defoliation. At the end of the drought period, all species showed increased defoliation and a related reduction in the concentration of sapwood soluble sugars. Defoliation was constrained by the BAI of the previous 5 years and sapwood production. No specific wood-inhabiting fungi were found in post-drought declining trees apart from blue-stain fungi, which extensively affected damaged Scots pines. Declining silver firs showed increases in BAI autocorrelation and variability prior to tree death. 4. Synthesis. Early warning signals of drought-triggered mortality seem to be species specific and reflect how different tree species cope with drought stress. Highly correlated declining growth patterns during drought can serve as a signal in silver fir, whereas changes in the content of sapwood soluble sugars are suitable vigour proxies for Scots and Aleppo pines. Longer growth and defoliation series, additional vigour parameters and multi-species comparisons are required to understand and predict drought-induced tree death.
Understanding how plants survive drought and cold is increasingly important as plants worldwide experience dieback with drought in moist places and grow taller with warming in cold ones. Crucial in plant climate adaptation are the diameters of water-transporting conduits. Sampling 537 species across climate zones dominated by angiosperms, we find that plant size is unambiguously the main driver of conduit diameter variation. And because taller plants have wider conduits, and wider conduits within species are more vulnerable to conduction-blocking embolisms, taller conspecifics should be more vulnerable than shorter ones, a prediction we confirm with a plantation experiment. As a result, maximum plant size should be short under drought and cold, which cause embolism, or increase if these pressures relax. That conduit diameter and embolism vulnerability are inseparably related to plant size helps explain why factors that interact with conduit diameter, such as drought or warming, are altering plant heights worldwide.
Forecasted increase drought frequency and severity may drive worldwide declines in forest productivity. Species-level responses to a drier world are likely to be influenced by their functional traits. Here, we analyse forest resilience to drought using an extensive network of tree-ring width data and satellite imagery. We compiled proxies of forest growth and productivity (TRWi, absolutely dated ring-width indices; NDVI, Normalized Difference Vegetation Index) for 11 tree species and 502 forests in Spain corresponding to Mediterranean, temperate, and continental biomes. Four different components of forest resilience to drought were calculated based on TRWi and NDVI data before, during, and after four major droughts (1986, 1994-1995, 1999, and 2005), and pointed out that TRWi data were more sensitive metrics of forest resilience to drought than NDVI data. Resilience was related to both drought severity and forest composition. Evergreen gymnosperms dominating semi-arid Mediterranean forests showed the lowest resistance to drought, but higher recovery than deciduous angiosperms dominating humid temperate forests. Moreover, semi-arid gymnosperm forests presented a negative temporal trend in the resistance to drought, but this pattern was absent in continental and temperate forests. Although gymnosperms in dry Mediterranean forests showed a faster recovery after drought, their recovery potential could be constrained if droughts become more frequent. Conversely, angiosperms and gymnosperms inhabiting temperate and continental sites might have problems to recover after more intense droughts since they resist drought but are less able to recover afterwards.
Aim The intensity and frequency of drought have increased considerably during recent decades in some Northern Hemisphere forested areas, and future climate warming could further magnify drought stress. We quantify how forests resist drought events and recover after them, i.e. we determine their growth resilience. Location North America and Europe. Methods We use a large tree‐ring database to study how drought influences forest growth resilience. We selected 775 tree‐ring width chronologies and studied the occurrence of years with extremely dry conditions (low soil moisture and/or high evaporative stress; hereafter ‘drought’) in these forests. For each drought in each forest we calculated three indices that represent different components of growth resilience to drought: resistance (Rt), recovery (Rc) and resilience (Rs). We related the variation in these indices with geographical, topographic, climatic and ecological conditions from each region. Results The three components of forest growth resilience were interrelated. Resistance and recovery were negatively related, and both were positively and nonlinearly related to resilience. Drought resistance increased with latitude, soil moisture and slope, whereas drought recovery decreased with latitude, soil moisture and summer normalized difference vegetation index. Drought resilience increased with elevation and decreased with the variation in soil moisture. Temperate broadleaf forests from wet regions showed a greater growth resistance (e.g. north‐eastern USA, central Europe) while conifer forests from dry to semi‐arid regions (e.g. south‐western USA, southern Europe) presented a greater growth recovery. Main conclusions The geographical patterns of growth resilience indices confirm the existence of different strategies among forests to cope with droughts, depending on the biome, the tree species and the prevailing climatic conditions. Geographical patterns in soil moisture availability tend to override species‐specific responses to drought.
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