Linking drought legacy effects across scales: From leaves to tree rings to ecosystems

Severe drought can cause lagged effects on tree physiology that negatively impact forest functioning for years. These “drought legacy effects” have been widely documented in tree‐ring records and could have important implications for our understanding of broader scale forest carbon cycling. However, legacy effects in tree‐ring increments may be decoupled from ecosystem fluxes due to (a) postdrought alterations in carbon allocation patterns; (b) temporal asynchrony between radial growth and carbon uptake; and (c) dendrochronological sampling biases. In order to link legacy effects from tree rings to whole forests, we leveraged a rich dataset from a Midwestern US forest that was severely impacted by a drought in 2012. At this site, we compiled tree‐ring records, leaf‐level gas exchange, eddy flux measurements, dendrometer band data, and satellite remote sensing estimates of greenness and leaf area before, during, and after the 2012 drought. After accounting for the relative abundance of tree species in the stand, we estimate that legacy effects led to ~10% reductions in tree‐ring width increments in the year following the severe drought. Despite this stand‐scale reduction in radial growth, we found that leaf‐level photosynthesis, gross primary productivity (GPP), and vegetation greenness were not suppressed in the year following the 2012 drought. Neither temporal asynchrony between radial growth and carbon uptake nor sampling biases could explain our observations of legacy effects in tree rings but not in GPP. Instead, elevated leaf‐level photosynthesis co‐occurred with reduced leaf area in early 2013, indicating that resources may have been allocated away from radial growth in conjunction with postdrought upregulation of photosynthesis and repair of canopy damage. Collectively, our results indicate that tree‐ring legacy effects were not observed in other canopy processes, and that postdrought canopy allocation could be an important mechanism that decouples tree‐ring signals from GPP.     

Drought legacies are dependent on water table depth,wood anatomy and drought timing across the eastern US

Severe droughts can impart long‐lasting legacies on forest ecosystems through lagged effects that hinder tree recovery and suppress whole‐forest carbon uptake. However, the local climatic and edaphic factors that interact to affect drought legacies in temperate forests remain unknown. Here, we pair a dataset of 143 tree ring chronologies across the mesic forests of the eastern US with historical climate and local soil properties. We found legacy effects to be widespread, the magnitude of which increased markedly in diffuse porous species, sites with deep water tables, and in response to late‐season droughts (August–September). Using an ensemble of downscaled climate projections, we additionally show that our sites are projected to drastically increase in water deficit and drought frequency by the end of the century, potentially increasing the size of legacy effects by up to 65% and acting as a significant process shaping forest composition, carbon uptake and mortality.     

Demographic shifts in eastern US forests increase the impact of late-season drought on forest growth

While forest communities are changing as a result of global environmental change, the impacts of tree species shifts on ecosystem services such as carbon storage are poorly quantified. In many parts of the eastern United States (US), more xeric-adapted oak-hickory dominated stands are being replaced with mesic beech-maple assemblages. To examine the possible impacts of this ongoing change in forest composition, we investigated how two wide-ranging and co-occurring eastern US species – Acer saccharum (sugar maple) and Quercus alba (white oak) – respond to interannual climate variability. Using 781 tree cores from 418 individual trees at 18 locations, we found late-growing season drought reduced A. saccharum growth more than that of Q. alba. A gradient in the growth reduction across latitude was also found in A. saccharum, where southern populations of A. saccharum experienced greater reductions in growth during drought. Drought had a legacy effect on growth for both species, with drought occurring later in the growing season having a larger legacy effect. Consequently, as forests shift from oak to maple dominance, drought in the later part of the growing season is likely to become an increasingly important control on forest productivity. Thus, our findings suggest that co-occurring species are responding to environmental conditions during different times in the growing season and, therefore, the timing of drought conditions will play an important role in forest productivity and carbon sequestration as forest species composition changes. These findings are particularly important because the projected increases in potential evapotranspiration, combined with possible changes in the seasonality of precipitation could have a substantial impact on how tree growth responds to future climatic change.     

Dendrochronological potential of four neotropical dry-forest tree species: Climate-growth correlations in northeast Brazil

Tropical dry forests (TDF) are highly important tropical forest ecosystems. Yet, these forests are highly threatened, usually neglected and only poorly studied. Understanding the long-term influences of environmental conditions on tree growth in these forests is crucial to understand the functioning, carbon dynamics and potential responses to future climate change of these forests. Dendrochronology can be used as a tool to provide these insights but has only scantly been applied in (dry) tropical forests. Here we evaluate the dendrochronological potential of four Caatinga neotropical dry forest tree species – Aspidosperma pyrifolium, Ziziphus joazeiro, Tabebuia aurea, and Libidibia ferrea – collected in two locations in northeastern Brazil (Sergipe state). We provide an anatomical characterization of the ring boundaries for the four species and investigate correlations of their growth with local and regional climatic variables. All four species form annual rings and show high inter-correlation (up to 0.806) and sensitivity (up to 0.565). Growth of all species correlated with local precipitation as well as with sea-surface temperatures in the tropical Atlantic and/or tropical Pacific oceans. We also show teleconnections between growth and the El Niño South Oscillation. The strong dependence of tree on precipitation is worrisome, considering that climate change scenarios forecast increased drought conditions in the Caatinga dry forest. Including more species and expanding dendrochronological studies to more areas would greatly improve our understanding of tree growth and functioning in TDFs. This type of knowledge is essential to assist the conservation, management and restoration of these critical tropical ecosystems.     

Legacies of more frequent drought in ponderosa pine across the western United States

Despite widespread interest in drought legacies—multiyear impacts of drought on tree growth—the key implication of reported drought legacies remains unaddressed: as impaired growth and slow recovery associated with drought legacies are pervasive across forest ecosystems, what is the impact of more frequent drought conditions? We investigated the assumption that either multiple drought years occurring during a short period (multiyear droughts), or droughts occurring during the recovery period from previous drought (compounded droughts), are detrimental to subsequent growth. There is evidence that drought responses may vary among populations of widespread species, leading us to examine regional differences in responses of the conifer Pinus ponderosa to historic drought frequency in the western United States. More frequent drought conditions incurred additional growth declines and shifts in growth–climate sensitivities in the years following drought relative to single‐drought events, with ‘triple‐droughts' being worse than ‘double‐droughts'. Notably, prediction skill was not strongly reduced when ignoring compounded droughts, a consequence of the temporally comprehensive formulation of our stochastic antecedent model that accounts for the climatic memory of tree growth. We argue that incorporating drought‐induced temporal variability in tree growth sensitivities can aid inference gained from statistical models, where more simplistic models could overestimate the severity of drought legacies. We also found regional differences in response to repeated drought, and suggest plastic post‐drought sensitivities and climatic memory may represent beneficial physiological adjustments in interior regions. Within‐species variability may thus mediate forest responses to increasing drought frequency under future climate change, but experimental approaches using more species are necessary to improve our understanding of the mechanisms that underlie drought legacy effects on tree growth.     

Overstorey–Understorey Interactions Intensify After Drought-Induced Forest Die-Off: Long-Term Effects for Forest Structure and Composition

Severe drought events increasingly affect forests worldwide, but little is known about their long-term effects at the ecosystem level. Competition between trees and herbs (‘overstorey–understorey competition’) for soil water can reduce tree growth and regeneration success and may thereby alter forest structure and composition. However, these effects are typically ignored in modelling studies. To test the long-term impact of water competition by the herbaceous understorey on forest dynamics, we incorporated this process in the dynamic forest landscape model LandClim. Simulations were performed both with and without understorey under current and future climate scenarios (RCP4.5 and RCP8.5) in a drought-prone inner-Alpine valley in Switzerland. Under current climate, herbaceous understorey reduced tree regeneration biomass by up to 51%, particularly in drought-prone landscape positions (i.e., south-facing, low-elevation slopes), where it also caused a shift in forest composition towards drought-tolerant tree species (for example, Quercus pubescens). For adult trees, the understorey had a minor effect on growth. Under future climate change scenarios, increasing drought frequency and intensity resulted in large-scale mortality of canopy trees, which intensified the competitive interaction between the understorey and tree regeneration. At the driest landscape positions, a complete exclusion of tree regeneration and a shift towards an open, savannah-like vegetation occurred. Overall, our results demonstrate that water competition by the herbaceous understorey can cause long-lasting legacy effects on forest structure and composition across drought-prone landscapes, by affecting the vulnerable recruitment phase. Ignoring herbaceous vegetation may thus lead to a strong underestimation of future drought impacts on forests.     

Climate-growth relations of congeneric tree species vary across a tropical vegetation gradient in Brazil

Seasonally dry tropical forests are an important global climatic regulator, a main driver of the global carbon sink dynamics and are predicted to suffer future reductions in their productivity due to climate change. Yet, little is known about how interannual climate variability affects tree growth and how climate-growth responses vary across rainfall gradients in these forests. Here we evaluate changes in climate sensitivity of tree growth along an environmental gradient of seasonally dry tropical vegetation types (evergreen forest – savannah – dry forest) in Northeastern Brazil, using congeneric species of two common neotropical genera: Aspidosperma and Handroanthus. We built tree-ring width chronologies for each species × forest type combinations and explored how growth variability correlated with local (precipitation, temperature) and global (the El Niño Southern Oscillation - ENSO) climatic factors. We also assessed how growth sensitivity to climate and the presence of growth deviations varied along the gradient. Precipitation stimulates tree growth and was the main growth-influencing factor across vegetation types. Trees in the dry forest site showed highest growth sensitivity to interannual variation in precipitation. Temperature and ENSO phenomena correlated negatively with growth and sensitivity to both climatic factors were similar across sites. Negative growth deviations were present and found mostly in the dry-forest species. Our results reveal a dominant effect of precipitation on tree growth in seasonally dry tropical forests and suggest that along the gradient, dry forests are the most sensitivity to drought. These forests may therefore be the most vulnerable to the deleterious effects of future climatic changes. These results highlight the importance of understanding the climatic sensitivity of different tropical forests. This understanding is key to predict the carbon dynamics in tropical regions, and sensitivity differences should be considered when prioritizing conservation measures of seasonally dry topical forests.     

Forest resilience to drought varies across biomes

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.     

Temperature and rainfall interact to control carbon cycling in tropical forests

Tropical forests dominate global terrestrial carbon (C) exchange, and recent droughts in the Amazon Basin have contributed to short‐term declines in terrestrial carbon dioxide uptake and storage. However, the effects of longer‐term climate variability on tropical forest carbon dynamics are still not well understood. We synthesised field data from more than 150 tropical forest sites to explore how climate regulates tropical forest aboveground net primary productivity (ANPP) and organic matter decomposition, and combined those data with two existing databases to explore climate – C relationships globally. While previous analyses have focused on the effects of either temperature or rainfall on ANPP, our results highlight the importance of interactions between temperature and rainfall on the C cycle. In cool forests (< 20 °C), high rainfall slowed rates of C cycling, but in warm tropical forests (> 20 °C) it consistently enhanced both ANPP and decomposition. At the global scale, our analysis showed an increase in ANPP with rainfall in relatively warm sites, inconsistent with declines in ANPP with rainfall reported previously. Overall, our results alter our understanding of climate – C cycle relationships, with high precipitation accelerating rates of C exchange with the atmosphere in the most productive biome on earth.     

Stable carbon isotope analysis reveals widespread drought stress in boreal black spruce forests

Unprecedented rates of climate warming over the past century have resulted in increased forest stress and mortality worldwide. Decreased tree growth in association with increasing temperatures is generally accepted as a signal of temperature‐induced drought stress. However, variations in tree growth alone do not reveal the physiological mechanisms behind recent changes in tree growth. Examining stable carbon isotope composition of tree rings in addition to tree growth can provide a secondary line of evidence for physiological drought stress. In this study, we examined patterns of black spruce growth and carbon isotopic composition in tree rings in response to climate warming and drying in the boreal forest of interior Alaska. We examined trees at three nested scales: landscape, toposequence, and a subsample of trees within the toposequence. At each scale, we studied the potential effects of differences in microclimate and moisture availability by sampling on northern and southern aspects. We found that black spruce radial growth responded negatively to monthly metrics of temperature at all examined scales, and we examined ?¹³C responses on a subsample of trees as representative of the wider region. The negative ?¹³C responses to temperature reveal that black spruce trees are experiencing moisture stress on both northern and southern aspects. Contrary to our expectations, ?¹³C from trees on the northern aspect exhibited the strongest drought signal. Our results highlight the prominence of drought stress in the boreal forest of interior Alaska. We conclude that if temperatures continue to warm, we can expect drought‐induced productivity declines across large regions of the boreal forest, even for trees located in cool and moist landscape positions.     

Amazon drought and its implications for forest flammability and tree growth: a basin-wide analysis

Severe drought in moist tropical forests provokes large carbon emissions by increasing forest flammability and tree mortality, and by suppressing tree growth. The frequency and severity of drought in the tropics may increase through stronger El Niño Southern Oscillation (ENSO) episodes, global warming, and rainfall inhibition by land use change. However, little is known about the spatial and temporal patterns of drought in moist tropical forests, and the complex relationships between patterns of drought and forest fire regimes, tree mortality, and productivity. We present a simple geographic information system soil water balance model, called RisQue (Risco de Queimada – Fire Risk) for the Amazon basin that we use to conduct an analysis of these patterns for 1996–2001. RisQue features a map of maximum plant‐available soil water (PAW_max) developed using 1565 soil texture profiles and empirical relationships between soil texture and critical soil water parameters. PAW is depleted by monthly evapotranspiration (ET) fields estimated using the Penman–Monteith equation and satellite‐derived radiation inputs and recharged by monthly rain fields estimated from 266 meteorological stations. Modeled PAW to 10 m depth (PAW_{10 m}) was similar to field measurements made in two Amazon forests. During the severe drought of 2001, PAW_{10 m} fell to below 25% of PAW_max in 31% of the region's forests and fell below 50% PAW_max in half of the forests. Field measurements and experimental forest fires indicate that soil moisture depletion below 25% PAW_max corresponds to a reduction in leaf area index of approximately 25%, increasing forest flammability. Hence, approximately one‐third of Amazon forests became susceptible to fire during the 2001 ENSO period. Field measurements also suggest that the ENSO drought of 2001 reduced carbon storage by approximately 0.2 Pg relative to years without severe soil moisture deficits. RisQue is sensitive to spin‐up time, rooting depth, and errors in ET estimates. Improvements in our ability to accurately model soil moisture content of Amazon forests will depend upon better understanding of forest rooting depths, which can extend to beyond 15 m. RisQue provides a tool for early detection of forest fire risk.     

Forest tree species adaptation to climate across biomes: Building on the legacy of ecological genetics to anticipate responses to climate change

Intraspecific variation plays a critical role in extant and future forest responses to climate change. Forest tree species with wide climatic niches rely on the intraspecific variation resulting from genetic adaptation and phenotypic plasticity to accommodate spatial and temporal climate variability. A centuries-old legacy of forest ecological genetics and provenance trials has provided a strong foundation upon which to continue building on this knowledge, which is critical to maintain climate-adapted forests. Our overall objective is to understand forest trees intraspecific responses to climate across species and biomes, while our specific objectives are to describe ecological genetics models used to build our foundational knowledge, summarize modeling approaches that have expanded the traditional toolset, and extensively review the literature from 1994 to 2021 to highlight the main contributions of this legacy and the new analyzes of provenance trials. We reviewed 103 studies comprising at least three common gardens, which covered 58 forest tree species, 28 of them with range-wide studies. Although studies using provenance trial data cover mostly commercially important forest tree species from temperate and boreal biomes, this synthesis provides a global overview of forest tree species adaptation to climate. We found that evidence for genetic adaptation to local climate is commonly present in the species studied (79%), being more common in conifers (87.5%) than in broadleaf species (67%). In 57% of the species, clines in fitness-related traits were associated with temperature variables, in 14% of the species with precipitation, and in 25% of the species with both. Evidence of adaptation lags was found in 50% of the species with range-wide studies. We conclude that ecological genetics models and analysis of provenance trial data provide excellent insights on intraspecific genetic variation, whereas the role and limits of phenotypic plasticity, which will likely determine the fate of extant forests, is vastly understudied.     

A comprehensive resilience assessment of Mexican tree species and their relationship with drought events over the last century

The resilience of forests to drought events has become a major natural resource sustainability concern, especially in response to climate change. Yet, little is known about the legacy effects of repeated droughts, and tree species ability to respond across environmental gradients. In this study, we used a tree-ring database (121 sites) to evaluate the overall resilience of tree species to drought events in the last century. We investigated how climate and geography affected the response at the species level. We evaluated temporal trends of resilience using a predictive mixed linear modeling approach. We found that pointer years (e.g., tree growth reduction) occurred during 11.3% of the 20th century, with an average decrease in tree growth of 66% compared to the previous period. The occurrence of pointer years was associated with negative values of the Standardized Precipitation Index (SPI, 81.6%) and Palmer Drought Severity Index (PDSI, 77.3%). Tree species differed in their resilience capacity, however, species inhabiting xeric conditions were less resistant but with higher recovery rates (e.g., Abies concolor, Pinus lambertiana, and Pinus jeffreyi). On average, tree species needed 2.7 years to recover from drought events, with extreme cases requiring more than a decade to reach pre-drought tree growth rates. The main abiotic factor related to resilience was precipitation, confirming that some tree species are better adapted to resist the effects of droughts. We found a temporal variation for all tree resilience indices (scaled to 100), with a decreasing resistance (−0.56 by decade) and resilience (−0.22 by decade), but with a higher recovery (+1.72 by decade) and relative resilience rate (+0.33 by decade). Our results emphasize the importance of time series of forest resilience, particularly by distinguishing the species-level response in the context of legacy of droughts, which are likely to become more frequent and intense under a changing climate.     

The impacts of increasing drought on forest dynamics,structure, and biodiversity in the United States

We synthesize insights from current understanding of drought impacts at stand‐to‐biogeographic scales, including management options, and we identify challenges to be addressed with new research. Large stand‐level shifts underway in western forests already are showing the importance of interactions involving drought, insects, and fire. Diebacks, changes in composition and structure, and shifting range limits are widely observed. In the eastern US, the effects of increasing drought are becoming better understood at the level of individual trees, but this knowledge cannot yet be confidently translated to predictions of changing structure and diversity of forest stands. While eastern forests have not experienced the types of changes seen in western forests in recent decades, they too are vulnerable to drought and could experience significant changes with increased severity, frequency, or duration in drought. Throughout the continental United States, the combination of projected large climate‐induced shifts in suitable habitat from modeling studies and limited potential for the rapid migration of tree populations suggests that changing tree and forest biogeography could substantially lag habitat shifts already underway. Forest management practices can partially ameliorate drought impacts through reductions in stand density, selection of drought‐tolerant species and genotypes, artificial regeneration, and the development of multistructured stands. However, silvicultural treatments also could exacerbate drought impacts unless implemented with careful attention to site and stand characteristics. Gaps in our understanding should motivate new research on the effects of interactions involving climate and other species at the stand scale and how interactions and multiple responses are represented in models. This assessment indicates that, without a stronger empirical basis for drought impacts at the stand scale, more complex models may provide limited guidance.     

Soil carbon dynamics in regrowing forest of eastern Amazonia

The future flora of Amazonia will include significant areas of secondary forest as degraded pastures are abandoned and secondary succession proceeds. The rate at which secondary forests regain carbon (C) stocks and re-establish biogeochemical cycles that resemble those of primary forests will influence the biogeochemistry of the region. Most studies have focused on the effects of deforestation on biogeochemical cycles. In this study, we present data on the recuperation of carbon stocks and carbon fluxes within a secondary forest of the eastern Amazon, and we compare these measurements to those for primary forest, degraded pasture, and productive pasture. Along a transect from a 23-y-old degraded pasture, through a 7-y-old secondary forest, through a 16-year-old secondary forest, and to a primary forest, the δ¹³C values of soil organic matter (SOM) in the top 10 cm of soil were – 21.0, – 26.5, – 27.4, and – 27.9‰, respectively, indicating that the isotopic signature of SOM from C3 forest plants was rapidly re-established. The degraded pasture also had significant inputs of C from C3 plants. Radiocarbon data indicated that most of the C in the top 10 cm of soil had been fixed by plants during the last 30 years. Differences in soil C inventory among land use types were small compared to uncertainties in their measurement. Root inputs were nearly identical in primary and secondary forests, and litterfall in the secondary forest was 88% of the litterfall rate of the primary forest. In contrast, the secondary forest had only 17% of the above ground biomass. Because of rapid cycling rates of soil C and rapid recovery of C fluxes to and from the soil, the below ground C cycle in this secondary forest was nearly identical with those of the unaltered primary forest.     

Components of tree resilience: effects of successive low‐growth episodes in old ponderosa pine forests

Recent world‐wide episodes of tree dieback have been attributed to increasing temperatures and associated drought. Because these events are likely to become more common, improved knowledge of their cumulative effects on resilience and the ability to recover pre‐disturbance conditions is important for forest management. Here we propose several indices to examine components of individual tree resilience based on tree ring growth: resistance (inverse of growth reduction during the episode), recovery (growth increase relative to the minimum growth during the episode), resilience (capacity to reach pre‐episode growth levels) and relative resilience (resilience weighted by the damage incurred during the episode). Based on tree ring analyses, we analyzed historical patterns of tree resilience to successive drought‐induced low growth periods in ponderosa pine trees growing in unmanaged, remote forests of the Rocky Mountains. Low‐growth periods registered in tree rings were related to anomalies in the Palmer drought severity index (PDSI) and were attributed to drought. Independently of the impact of a specific event, subsequent growth after a single low‐growth episode was related to the growth prior to the event. Growth performance differed with tree age: young trees were overall more resistant to low‐growth periods, but older trees recovered better from more recent events. Regardless of tree age, recently burned sites exhibited lower post‐episode growth and lower resistance and resilience than unburned ones. We found mixed evidence for the cumulative effect of past low‐growth episodes: overall, greater impacts of a prior event and greater cumulative effects of past low‐growth periods caused a decrease in resistance. However, we did not find a progressive decrease in resilience over time in old trees. Our results highlight the value of using a combination of estimators to evaluate the different components of resilience. Specifically, while tree responses to disturbance depend on past disturbance episodes, the response is context‐specific and depends on the impact the capacity to recover after disturbance. This suggests that recent increases in forest mortality under current climate trends could relate to thresholds on specific components of resilience (resistance, recovery, resilience itself) rather than to an overall loss of resilience over time. Identifying such thresholds and their underlying mechanisms is a promising area of research with important implications for forest management.     

菌根真菌影响森林生态系统碳循环研究进展

森林生态系统在全球碳(C)储量中占据极为重要的地位。菌根真菌广泛存在于森林生态系统中,在森林生态系统C循环过程中发挥重要的作用。阐述了不同菌根类型真菌在森林生态系统C循环过程中的功能,对比了温带/北方森林与热带/亚热带森林中菌根真菌介导的C循环研究方面新近取得的研究结果。发现温带和北方森林的外生菌根(EcM)植物对地上生物量C的贡献相对较小,然而是地下C储量的主要贡献者;以丛枝菌根(AM)共生为主的热带/亚热带森林地表生物量占比较高,表明AM植被对热带/亚热带森林地上生物量C的贡献相对较大。我们还就全球变化背景下,菌根真菌及其介导的森林生态系统C汇功能,以及不同菌根类型树种影响C循环的机制等进行了总结。菌根真菌通过影响凋落物分解、土壤有机质形成及地下根系生物量,进而影响整个森林生态系统的C循环功能。菌根介导的森林C循环过程很大程度上取决于(优势)树木的菌根类型和森林土壤中菌根真菌的群落结构。最后指出了当前研究存在的主要问题以及未来研究展望。本文旨在明确菌根真菌在森林生态系统C循环转化过程中的重要生态功能,有助于准确地评估森林生态系统C汇现状,为应对全球变化等提供重要的依据。     

A first assessment of the impact of the extreme 2018 summer drought on Central European forests

In 2018, Central Europe experienced one of the most severe and long-lasting summer drought and heat wave ever recorded. Before 2018, the 2003 millennial drought was often invoked as the example of a “hotter drought”, and was classified as the most severe event in Europe for the last 500 years. First insights now confirm that the 2018 drought event was climatically more extreme and had a greater impact on forest ecosystems of Austria, Germany and Switzerland than the 2003 drought. Across this region, mean growing season air temperature from April to October was more than 3.3°C above the long-term average, and 1.2°C warmer than in 2003. Here, we present a first impact assessment of the severe 2018 summer drought and heatwave on Central European forests. In response to the 2018 event, most ecologically and economically important tree species in temperate forests of Austria, Germany and Switzerland showed severe signs of drought stress. These symptoms included exceptionally low foliar water potentials crossing the threshold for xylem hydraulic failure in many species and observations of widespread leaf discoloration and premature leaf shedding. As a result of the extreme drought stress, the 2018 event caused unprecedented drought-induced tree mortality in many species throughout the region. Moreover, unexpectedly strong drought-legacy effects were detected in 2019. This implies that the physiological recovery of trees was impaired after the 2018 drought event, leaving them highly vulnerable to secondary drought impacts such as insect or fungal pathogen attacks. As a consequence, mortality of trees triggered by the 2018 events is likely to continue for several years. Our assessment indicates that many common temperate European forest tree species are more vulnerable to extreme summer drought and heat waves than previously thought. As drought and heat events are likely to occur more frequently with the progression of climate change, temperate European forests might approach the point for a substantial ecological and economic transition. Our assessment also highlights the urgent need for a pan-European ground-based monitoring network suited to track individual tree mortality, supported by remote sensing products with high spatial and temporal resolution to track, analyse and forecast these transitions.     

Immediate and carry-over effects of late-spring frost and growing season drought on forest gross primary productivity capacity in the Northern Hemisphere

Forests are increasingly exposed to extreme global warming-induced climatic events. However, the immediate and carry-over effects of extreme events on forests are still poorly understood. Gross primary productivity (GPP) capacity is regarded as a good proxy of the ecosystem's functional stability, reflecting its physiological response to its surroundings. Using eddy covariance data from 34 forest sites in the Northern Hemisphere, we analyzed the immediate and carry-over effects of late-spring frost (LSF) and growing season drought on needle-leaf and broadleaf forests. Path analysis was applied to reveal the plausible reasons behind the varied responses of forests to extreme events. The results show that LSF had clear immediate effects on the GPP capacity of both needle-leaf and broadleaf forests. However, GPP capacity in needle-leaf forests was more sensitive to drought than in broadleaf forests. There was no interaction between LSF and drought in either needle-leaf or broadleaf forests. Drought effects were still visible when LSF and drought coexisted in needle-leaf forests. Path analysis further showed that the response of GPP capacity to drought differed between needle-leaf and broadleaf forests, mainly due to the difference in the sensitivity of canopy conductance. Moreover, LSF had a more severe and long-lasting carry-over effect on forests than drought. These results enrich our understanding of the mechanisms of forest response to extreme events across forest types.     

Patterns of drought tolerance in major European temperate forest trees: climatic drivers and levels of variability

The future performance of native tree species under climate change conditions is frequently discussed, since increasingly severe and more frequent drought events are expected to become a major risk for forest ecosystems. To improve our understanding of the drought tolerance of the three common European temperate forest tree species Norway spruce, silver fir and common beech, we tested the influence of climate and tree‐specific traits on the inter and intrasite variability in drought responses of these species. Basal area increment data from a large tree‐ring network in Southern Germany and Alpine Austria along a climatic cline from warm‐dry to cool‐wet conditions were used to calculate indices of tolerance to drought events and their variability at the level of individual trees and populations. General patterns of tolerance indicated a high vulnerability of Norway spruce in comparison to fir and beech and a strong influence of bioclimatic conditions on drought response for all species. On the level of individual trees, low‐growth rates prior to drought events, high competitive status and low age favored resilience in growth response to drought. Consequently, drought events led to heterogeneous and variable response patterns in forests stands. These findings may support the idea of deliberately using spontaneous selection and adaption effects as a passive strategy of forest management under climate change conditions, especially a strong directional selection for more tolerant individuals when frequency and intensity of summer droughts will increase in the course of global climate change.