1. Climate models predict increasing drought intensity and frequency for many regions, which may have negative consequences for tree recruitment, growth and mortality, as well as forest ecosystem services. Furthermore, practical strategies for minimizing vulnerability to drought are limited. Tree population density, a metric of tree abundance in a given area, is a primary driver of competitive intensity among trees, which influences tree growth and mortality. Manipulating tree population density may be a mechanism for moderating drought-induced stress and growth reductions, although the relationship between tree population density and tree drought vulnerability remains poorly quantified, especially across climatic gradients. 2. In this study, we examined three long-term forest ecosystem experiments in two widely distributed North American pine species, ponderosa pine Pinus ponderosa (Lawson & C. Lawson) and red pine Pinus resinosa (Aiton), to better elucidate the relationship between tree population density, growth and drought. These experiments span a broad latitude and aridity range and include tree population density treatments that have been purposefully maintained for several decades. We investigated how tree population density influenced resistance (growth during drought) and resilience (growth after drought compared to pre-drought growth) of stand-level growth during and after documented drought events. 3. Our results show that relative tree population density was negatively related to drought resistance and resilience, indicating that trees growing at lower densities were less vulnerable to drought. This result was apparent in all three forest ecosystems, and was consistent across species, stand age and drought intensity. 4. Synthesis and applications. Our results highlighted that managing pine forest ecosystems at low tree population density represents a promising adaptive strategy for reducing the adverse impacts of drought on forest growth in coming decades. Nonetheless, the broader applicability of our findings to other types of forest ecosystems merits additional investigation.
Extreme climate events (ECEs) such as severe droughts, heat waves, and late spring frosts are rare but exert a paramount role in shaping tree species distributions. The frequency of such ECEs is expected to increase with climate warming, threatening the sustainability of temperate forests. Here, we analyzed 2,844 tree‐ring width series of five dominant European tree species from 104 Swiss sites ranging from 400 to 2,200 m a.s.l. for the period 1930–2016. We found that (a) the broadleaved oak and beech are sensitive to late frosts that strongly reduce current year growth; however, tree growth is highly resilient and fully recovers within 2 years; (b) radial growth of the conifers larch and spruce is strongly and enduringly reduced by spring droughts—these species are the least resistant and resilient to droughts; (c) oak, silver fir, and to a lower extent beech, show higher resistance and resilience to spring droughts and seem therefore better adapted to the future climate. Our results allow a robust comparison of the tree growth responses to drought and spring frost across large climatic gradients and provide striking evidence that the growth of some of the most abundant and economically important European tree species will be increasingly limited by climate warming. These results could serve for supporting species selection to maintain the sustainability of forest ecosystem services under the expected increase in ECEs.
Forests around the world are experiencing increasingly severe droughts and elevated competitive intensity due to increased tree density. However, the influence of interactions between drought and competition on forest growth remains poorly understood. Using a unique dataset of stand‐scale dendrochronology sampled from 6405 trees, we quantified how annual growth of entire tree populations responds to drought and competition in eight, long‐term (multi‐decadal), experiments with replicated levels of density (e.g., competitive intensity) arrayed across a broad climatic and compositional gradient. Forest growth (cumulative individual tree growth within a stand) declined during drought, especially during more severe drought in drier climates. Forest growth declines were exacerbated by high density at all sites but one, particularly during periods of more severe drought. Surprisingly, the influence of forest density was persistent overall, but these density impacts were greater in the humid sites than in more arid sites. Significant density impacts occurred during periods of more extreme drought, and during warmer temperatures in the semi‐arid sites but during periods of cooler temperatures in the humid sites. Because competition has a consistent influence over growth response to drought, maintaining forests at lower density may enhance resilience to drought in all climates.
Global climate change is expected to further raise the frequency and severity of extreme events, such as droughts. The effects of extreme droughts on trees are difficult to disentangle given the inherent complexity of drought events (frequency, severity, duration, and timing during the growing season). Besides, drought effects might be modulated by trees’ phenotypic variability, which is, in turn, affected by long‐term local selective pressures and management legacies. Here we investigated the magnitude and the temporal changes of tree‐level resilience (i.e., resistance, recovery, and resilience) to extreme droughts. Moreover, we assessed the tree‐, site‐, and drought‐related factors and their interactions driving the tree‐level resilience to extreme droughts. We used a tree‐ring network of the widely distributed Scots pine ( Pinus sylvestris ) along a 2,800 km latitudinal gradient from southern Spain to northern Germany. We found that the resilience to extreme drought decreased in mid‐elevation and low productivity sites from 1980–1999 to 2000–2011 likely due to more frequent and severe droughts in the later period. Our study showed that the impact of drought on tree‐level resilience was not dependent on its latitudinal location, but rather on the type of sites trees were growing at and on their growth performances (i.e., magnitude and variability of growth) during the predrought period. We found significant interactive effects between drought duration and tree growth prior to drought, suggesting that Scots pine trees with higher magnitude and variability of growth in the long term are more vulnerable to long and severe droughts. Moreover, our results indicate that Scots pine trees that experienced more frequent droughts over the long‐term were less resistant to extreme droughts. We, therefore, conclude that the physiological resilience to extreme droughts might be constrained by their growth prior to drought, and that more frequent and longer drought periods may overstrain their potential for acclimation.
Climate-induced tree mortality became a global phenomenon during the last century and it is expected to increase in many regions in the future along with a further increase in the frequency of drought and heat events. However, tree mortality at the ecosystem level remains challenging to quantify since long-term, tree-individual, reliable observations are scarce. Here, we present a unique data set of monitoring records from 276 permanent plots located in 95 forest stands across Switzerland, which include five major European tree species (Norway spruce, Scots pine, silver fir, European beech, and sessile and common oak) and cover a time span of over one century (1898–2013), with inventory periods of 5–10 years. The long-term average annual mortality rate of the investigated forest stands was 1.5%. In general, species-specific annual mortality rates did not consistently increase over the last decades, except for Scots pine forests at lower altitudes, which exhibited a clear increase of mortality since the 1960s. Temporal trends of tree mortality varied also depending on diameter at breast height (DBH), with large trees generally experiencing an increase in mortality, while mortality of small trees tended to decrease. Normalized mortality rates were remarkably similar between species and a modest, but a consistent and steady increasing trend was apparent throughout the study period. Mixed effects models revealed that gradually changing stand parameters (stand basal area and stand age) had the strongest impact on mortality rates, modulated by climate, which had increasing importance during the last decades. Hereby, recent climatic changes had highly variable effects on tree mortality rates, depending on the species in combination with abiotic and biotic stand and site conditions. This suggests that forest species composition and species ranges may change under future climate conditions. Our data set highlights the complexity of forest dynamical processes such as long-term, gradual changes of forest structure, demography and species composition, which together with climate determine mortality rates.
Foresters from many countries are seeking for tree species or provenances able to cope with expected climate change. While it becomes clear that some temperate tree species will increasingly suffer from climate warming, the fate of the ecologically and economically important silver fir (Abies alba Mill.) remains uncertain and debated because the ecological requirements of this species, as well as its resilience to drought are still unclear. On the one hand, paleoecological studies reveal that this species was widely distributed under much warmer climate, suggesting a high potential to cope with ongoing and future climate warming. On the other hand, species distribution models (SDMs) generally predict a strong decline of its climatic suitability in the future. This paper aims to clarify the potential of this species to thrive in central and western Europe under predicted climate warming by reviewing the knowledge gained from different fields. Based on insight from different fields, we argue that silver fir has a great potential to thrive under warmer conditions in western and central Europe provided sufficient rainfall, as forecasted by climate models for most regions by 2100. For instance, dendroecological studies demonstrate that silver fir is more resistant and resilient to drought compared to co-occurring species such as Norway spruce, European beech and larch. The most prominent obstacle for increasing the proportion of fir in mixed forests nowadays is ungulate browsing that often prevents its upgrowth.
We investigated forest canopy gaps in the mixed beech (Fagus sylvatica L.), silver fir (Abies alba Miller), and Norway spruce (Picea abies (L.) Karst.) old-growth forest of Lom in the Dinaric Mountains of Bosnia and Herzegovina. Gap size, age, gap fraction, gapmaker characteristics and the structure and composition of gapfillers were documented to investigate gap dynamics. The percentages of forest area in canopy and expanded gaps were 19% and 41%, respectively. The median canopy gap size was 77 m 2 , and ranged from 11 to 708 m 2. Although there were many single tree-fall gaps, the majority had multiple gapmakers that were often in different stages of decay, suggesting gap expansion is important at the study site. Of the gapmakers recorded, 14% were uprooted stems, 60% snapped stems, and 26% were standing dead trees. Dendroecological analysis suggests that gap formation varied in time. The density of gapfillers was not correlated to gap size, and the species composition of gapfillers varied between seedling, sapling, and tree life stages. The results suggest that gaps are mainly formed by endogenous senescence of single canopy trees. Exogenous disturbance agents, most likely related to wind and snow, act mainly as secondary agents in breaking weakened trees and in expanding previously established gaps. Although the findings are partially consistent with other studies of gap disturbance processes in similar old-growth forests in central Europe, the observed gap dynamic places the Lom core area at the end of a gradient that ranges from forests controlled by very small-scale processes to those where large, stand replacing disturbances predominate.
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