Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks.

Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, >3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality.

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.

Mixed species forests can often be more productive and deliver higher levels of ecosystem services and functions than monocultures. However, complementarity effects for any given tree species are difficult to generalize because they can vary greatly along gradients of climatic conditions and resource availability. Identifying the conditions where species diversity can positively influence productivity is crucial. To date, few studies have examined how growth complementarity across species and mixture types is modulated by stand and environmental factors, and fewer have considered more than one or two factors. We investigated how complementarity effects for several major Central European tree species change with climatic and edaphic conditions, and with stand structural characteristics, including species composition. We used data from the Swiss National Forest Inventory, which is based on 3,231 plots of pure and mixed stands (19 mixture types) across a broad environmental gradient, to test (i) how mixing effects change depending on the identity of the admixed species and (ii) if complementarity consistently increases when environmental conditions become harsher. The magnitude, whether positive or negative, of complementarity increased with increasing stand density and stand developmental stage, but no general pattern could be identified across mixture types. Complementarity for many species increased as drought intensity and temperature increased, but not for all species and mixture types. While soil conditions, nitrogen and site topography influenced complementarity for many species, there was no general pattern (increases and decreases were observed). Synthesis. Our study indicates that complementarity varies strongly with stand density and stand development as well as with topographic, climatic and soil conditions. This emphasizes the need to account for site‐dependent conditions when exploring mixture effects in relation to forest productivity. We found that under certain conditions (i.e. increasing drought, higher temperature), mixed forests can promote individual tree growth in Central European temperate forests. However, careful assessments depending on the species composing the stands are required under changing resource availability as well as under different levels of stand density and development.

Tree mortality is a key driver of forest dynamics and its occurrence is projected to increase in the future due to climate change. Despite recent advances in our understanding of the physiological mechanisms leading to death, we still lack robust indicators of mortality risk that could be applied at the individual tree scale. Here, we build on a previous contribution exploring the differences in growth level between trees that died and survived a given mortality event to assess whether changes in temporal autocorrelation, variance, and synchrony in time-series of annual radial growth data can be used as early warning signals of mortality risk. Taking advantage of a unique global ring-width database of 3065 dead trees and 4389 living trees growing together at 198 sites (belonging to 36 gymnosperm and angiosperm species), we analyzed temporal changes in autocorrelation, variance, and synchrony before tree death (diachronic analysis), and also compared these metrics between trees that died and trees that survived a given mortality event (synchronic analysis). Changes in autocorrelation were a poor indicator of mortality risk. However, we found a gradual increase in inter-annual growth variability and a decrease in growth synchrony in the last ∼20 years before mortality of gymnosperms, irrespective of the cause of mortality. These changes could be associated with drought-induced alterations in carbon economy and allocation patterns. In angiosperms, we did not find any consistent changes in any metric. Such lack of any signal might be explained by the relatively high capacity of angiosperms to recover after a stress-induced growth decline. Our analysis provides a robust method for estimating early-warning signals of tree mortality based on annual growth data. In addition to the frequently reported decrease in growth rates, an increase in inter-annual growth variability and a decrease in growth synchrony may be powerful predictors of gymnosperm mortality risk, but not necessarily so for angiosperms.

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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.

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