Tree growth response to drought and temperature in a mountain landscape in northern Arizona, USA

Aim To understand how tree growth response to regional drought and temperature varies between tree species, elevations and forest types in a mountain landscape. Location Twenty‐one sites on an elevation gradient of 1500 m on the San Francisco Peaks, northern Arizona, USA. Methods Tree‐ring data for the years 1950–2000 for eight tree species (Abies lasiocarpa var. arizonica (Merriam) Lemm., Picea engelmannii Parry ex Engelm., Pinus aristata Engelm., Pinus edulis Engelm., Pinus flexilis James, Pinus ponderosa Dougl. ex Laws., Pseudotsuga menziesii var. glauca (Beissn.) Franco and Quercus gambelii Nutt.) were used to compare sensitivity of radial growth to regional drought and temperature among co‐occurring species at the same site, and between sites that differed in elevation and species composition. Results For Picea engelmannii, Pinus flexilis, Pinus ponderosa and Pseudotsuga menziesii, trees in drier, low‐elevation stands generally had greater sensitivity of radial growth to regional drought than trees of the same species in wetter, high‐elevation stands. Species low in their elevational range had greater drought sensitivity than co‐occurring species high in their elevational range at the pinyon‐juniper/ponderosa pine forest ecotone, ponderosa pine/mixed conifer forest ecotone and high‐elevation invaded meadows, but not at the mixed conifer/subalpine forest ecotone. Sensitivity of radial growth to regional drought was greater at drier, low‐elevation compared with wetter, high‐elevation forests. Yearly growth was positively correlated with measures of regional water availability at all sites, except high‐elevation invaded meadows where growth was weakly correlated with all climatic factors. Yearly growth in high‐elevation forests up to 3300 m a.s.l. was more strongly correlated with water availability than temperature. Main conclusions Severe regional drought reduced growth of all dominant tree species over a gradient of precipitation and temperature represented by a 1500‐m change in elevation, but response to drought varied between species and stands. Growth was reduced the most in drier, low‐elevation forests and in species growing low in their elevational range in ecotones, and the least for trees that had recently invaded high‐elevation meadows. Constraints on tree growth from drought and high temperature are important for high‐elevation subalpine forests located near the southern‐most range of the dominant species.     

Resilience capacity of Araucaria araucana to extreme drought events

Ongoing climate change has induced modification in the frequency and intensity of extreme climatic events, with consequent impact on tree and forest growth resilience. Araucaria araucana is an endangered Patagonian conifer, which provides several ecosystem services to local human societies and plays fundamental ecological roles in natural communities. These woodlands have historically suffered different types of anthropogenic disturbance, such as fire, logging and grazing, nevertheless the species resilience to extreme drought events remains still poorly understood. To fill this gap of knowledge, we applied dendrochronological methods to several A. araucana stands distributed along a steep bioclimatic gradient in order to reconstruct resilience capacity, in term of stem growth resistance and recovery, to three successive extreme spring-early summer droughts which occurred during the 20th century. Results showed an increase in the species recovery along the considered dry spells, whereas no clear trend emerged for resistance, suggesting no cumulative effect of drought upon resilience. Both resistance and recovery presented different values depending on bioclimatic settings, being xeric stands more sensitive to extreme episodes with respect to mesic woodlands, particularly during the more recent drought event when trees growing in drier environments were not able to reach pre-drought stem growth rates. Tree-level characteristics, such as age and growth trends prior to drought, modulated the species resilience, suggesting that future dry spells would possibly induce shifts in population dynamics, and furthermore be detrimental for fast-growing trees. Our analysis highlighted the response of a key Patagonian tree species to extreme drought events, providing bioclimatic-specific useful information for conservation plans of this natural resource.     

西藏东部主要建群树种径向生长对极端干旱的响应差异

不同坡向、不同海拔树木生长对极端干旱事件的响应可能不同,然而这方面的认识不足。为此,选取西藏东部珠角拉山阴、阳坡的建群树种川西云杉 (Picea likiangensis var. rubescens) 和大果圆柏 (Juniperus tibetica),在不同海拔高度建立了树轮宽度年表,分析了径向生长的气候响应,以及对极端干旱事件的抵抗力和恢复力。结果表明:阳坡大果圆柏和阴坡川西云杉的树木生长对气候的响应存在相似性,均与前一年3-6月、11-12月气温显著正相关,与当年4-5月气温显著负相关,与当年4-5月降水和帕尔默干旱指数 (PDSI, Palmer Drought Severity Index) 显著正相关 (P <0.05)。阳坡大果圆柏的抵抗力显著低于阴坡川西云杉。随着海拔升高,阴坡川西云杉树木个体的抵抗力和恢复力均显著提高,而阳坡大果圆柏树木个体抵抗力、恢复力在不同海拔无显著区别。结合混合效应模型表明树木抵抗力主要受当年4-5月平均最高气温限制,树木恢复力主要受干旱事件后四年4-5月平均最高气温限制 (P <0.01),说明生长季高温引起的极端干旱是树木径向生长下降的主要原因。     

Abiotic factors modulate post-drought growth resilience of Scots pine plantations and rear-edge Scots pine and oak forests

The proportion of planted forests in the Mediterranean Basin is one of the largest in the world. These plantations are dominated by pine species and present a series of characteristics such as low elevation, high competition or small tree size that make them more vulnerable to droughts. However, quantitative assessments of their post-drought growth resilience in accordance with species, site factors and tree characteristics are lacking. In this study we sampled 164 trees at four forest sites located in the drought-prone Sierra Nevada, southeastern Spain. We compared growth responsiveness to drought in rear-edge planted vs. relic natural Scots pine (Pinus sylvestris) and coexisting Pyrenean oak (Quercus pyrenaica) stands. Our objective was to characterize and compare the different growth responses to drought between species and sites and the effect of the main physiographic factors (altitude, aspect, and slope) on these responses since the influence of these factors on post-drought resistance and resilience has received little attention to date. Our results reveal that the planted pine sites with the lowest mean growth rates displayed greater resistance during drought, and that higher altitude was associated with improved resistance and/or resilience for all species and sites. Natural pine and Pyrenean oak stands were better adapted to the dry climatic conditions of the Mediterranean region where the study was undertaken, displaying greater resistance and/or resilience and lower influence of drought on growth in comparison to stands of planted pines. These results suggest that promoting the conservation of high-elevation pine plantations and enhancing the regeneration of natural pine and oak may improve the resistance and resilience of these drought-prone forest ecosystems.     

Austrocedrus chilensis growth decline in relation to drought events in northern Patagonia, Argentina

The significant mortality of the Austrocedrus chilensis (D. Don) Pic. Serm. et Bizarri forests, locally known as “Mal del Ciprés”, has been reported since 1945 for most sites across its distribution in Argentina. However, the cause of this decline is still a topic of discussion. In this study, radial growth patterns from symptomatic and asymptomatic A. chilensis trees were analyzed to determine the influence of drought events on tree growth. Fifty pairs of symptomatic and asymptomatic trees with similar DBH, competition, and microsite conditions were cored at five pure A. chilensis stands near El Bolsón, Río Negro, Argentina. A reference chronology from nonaffected trees was used to cross-date all cores and to determine the relationship between A. chilensis radial growth and climate. The growth of A. chilensis is favored by above average precipitation in late spring–early summer (November and December). A strong relationship was also observed between radial growth patterns and the Palmer drought severity index, a measure of the regional water deficit. Significant differences in growth patterns were recorded between symptomatic and asymptomatic trees. Following extreme drought events, the growth of symptomatic trees is consistently lower than in asymptomatic trees. Based on the larger number of droughts recorded during the past decades and on future climatic predictions suggesting increasing trends in the frequency and intensity of drought events in northern Patagonia, a gradual increase in the number of trees affected by “Mal del Ciprés” along the twenty-first century is likely expected.     

Contrasting climate influences on Nothofagus pumilio establishment along elevational gradients

Predicted warmer temperatures and more frequent extreme climatic events in the southern Andes may affect the dynamics of the Patagonian forests. These environmental changes may differentially alter the probability of Nothofagus pumilio establishment across its altitudinal range of distribution. We monitored fruit fall, seedling emergence and survival at altitudinal distribution range of N. pumilio forests in Santa Cruz (49°22′ S—72°56′ W), Argentina. Fruit fall, seedling emergence and survival were tested in relation to drought, based on the SPEI (Standardized Precipitation Evapotranspiration Index), interacting with elevation. Fruit fall was consistently higher at low elevation and the abundance of fruits was not affected by drought intensity. Density of new-born seedlings was?~?9 to 24 times lower at low- than at high-elevation plots in our first observations (2014 and 2016), characterized by warm-dry climatic conditions in spring-early summers. In contrast, seedling abundance was?~?1.5 times larger at low elevations during relatively cold-wet growing seasons. Survival probability was explained by the interaction between SPEI and elevation. At low elevation and in dry periods, survival probability was lower (CI 54–72%) than in wet periods (CI 68–84%) but at high elevation similar survival was registered even with positive or negative SPEI values. Our results show interacting effects of elevation and drought on tree establishment at the elevation limits, with positive and negative drought effects at high and low elevations, respectively. Predicted increase of extreme drought events during the XXI century could be detrimental for N. pumilio establishment at dry, low-elevation forests.

     

Size‐dependence of tree growth response to drought for Norway spruce and European beech individuals in monospecific and mixed‐species stands

• Climate anomalies have resulted in changing forest productivity, increasing tree mortality in Central and Southern Europe. This has resulted in more severe and frequent ecological disturbances to forest stands. This study analysed the size‐dependence of growth response to drought years based on 384 tree individuals of Norway spruce [Picea abies (L.) Karst.] and European beech [Fagus sylvatica ([L.)] in Bavaria, Germany.
• Samples were collected in both monospecific and mixed‐species stands. To quantify the growth response to drought stress, indices for basal area increment, resistance, recovery and resilience were calculated from tree ring measurements of increment cores. Linear mixed models were developed to estimate the influence of drought periods.
• The results show that ageing‐related growth decline is significant in drought years. Drought resilience and resistance decrease significantly with growth size among Norway spruce individuals. Evidence is also provided for robustness in the resilience capacity of European beech during drought stress. Spruce benefits from species mixing with deciduous beech, with over‐yielding spruce in pure stands.
• The importance of the influence of size‐dependence within tree growth studies during disturbances is highlighted and should be considered in future studies of disturbances, including drought.

     

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.     

秦岭中西部油松径向生长对气候因子的响应差异研究

在气候变暖背景下,树木径向生长对气候因子的响应会随区域干湿变化而有所差异。秦岭属于气候敏感区和生态脆弱区,南北气候特征差异明显,分析气候变化背景下树木径向生长对气候因子及干旱事件的响应,对准确预测未来气候变化对树木生长的影响至关重要。为探究该地区不同干湿环境下油松对气候因子及干旱事件响应的特点及差异,共采集秦岭中西部南北坡共4个样点的油松树轮样芯,利用树木年轮生态学的方法,分析各地油松年表与气候因子之间的关系,通过计算抵抗力、恢复力、恢复弹力等指标探究树木径向生长对干旱事件的应对能力,结果表明：1)在西部和北坡的3个样点,油松径向生长主要与前一年7—9月、当年5—7月的气温呈显著负相关,与当年5月降水呈显著正相关,在中部南坡油松径向生长主要与当年2—4月、9月气温呈显著正相关,与当年4月降水呈显著负相关;2)生长季气温升高所引发的水分亏缺,是研究区西部和北坡油松径向生长受限制的主要原因,且中部南坡油松生长受气温和干旱因子的制约要明显弱于其它3个样点;3)西部北坡油松对干旱的抵抗能力、恢复能力及恢复弹力均弱于西部南坡及中部北坡油松。在未来对研究区树木的管理与保护工作中,应更加关注西部北坡...     

Impact of extreme drought on tree-ring width and vessel anatomical features of Chukrasia tabularis

Multiple sources of evidence suggest an increasing frequency of extreme climatic events during the past century. In Bangladesh, a country strongly influenced by the South Asian monsoon climate, the years 1999 and 2006 were the most severe droughts among the ten drought events identified over the last four decades. We investigated the impact of these two drought events on radial growth and xylem anatomical features of the brevi-deciduous tree species Chukrasia tabularis in a moist tropical forest in Bangladesh. Tree radial growth declined by 54% during the 1999 and 48.7% during the 2006 droughts, respectively. Among the wood anatomical features, the number of vessels (NV) showed the highest sensitivity to drought, with a 45% decrease in the 1999 drought year, followed by total vessel area (TVA) and mean vessel area (MVA). On the other hand, Vessel density (VD) increased by 13% during the 1999 drought but the increase in VD was very low in the drought year 2006. The decreasing vessel area and increasing vessel density indicate xylem hydraulic adaptation of C. tabularis to minimize drought induced cavitation risk and to avoid hydraulic failure. The significant correlations between the Standardized Precipitation Evapotranspiration Index (SPEI) and time series of tree-ring width and vessel variables imply that decline in radial growth and changes in vessel features in C. tabularis are likely to be caused by drought induced water stress. Our analyses suggest that radial growth and wood anatomical features of C. tabularis are highly sensitive to extreme drought events in South Asian moist tropical forests and can be used to reconstruct past droughts and to model tree response to drought stress under future climate conditions.     

山西忻州地区1900-2012年典型森林的健康历史

健康的森林对维持其区域生态系统服务起着至关重要的作用,了解树木的生长历史对评估气候变化背景下森林的健康状况具有促进作用。选择山西高原中北部忻州地区保存较好的4个森林为研究对象,利用树木年轮学方法分析其生长变化特征,获得了该地区森林健康的时间和空间特征。结果表明：该地区森林在过去1个世纪中,存在3次不健康事件（1910-1940、1970-1987和1990-2012年）且在空间上表现出明显的同步性,不同时期的不健康事件持续时间和强度不尽相同,1930s的不健康事件持续时间最长也最为显着。树木径向生长与气象观测资料的相关分析显示,该地区森林生长主要与当年5-6月份温度呈负相关,与7月份降水和5-9月PDSI指数显着正相关。生长季持续的高温或降水减少造成的极端干旱事件是不健康事件空间一致性的主要驱动力,各采样点树种以及林分组成差异是影响时间特征不一致的可能原因。研究结果提供的森林健康历史数据对评估极端气候条件对森林健康生长的影响及制定合理的森林保护措施具有积极的现实意义。     

Dendrochronology in Coniferous forests around Lake Rara,West Nepal

In two forest stands, one domonated byAbies spectabilis, and the other byPinus wallichiana-Picea smithiana, 198 cores were taken from 105 conifers in May 1983 and the annual ring widths were measured. The annual ring widths usually had significant similarities between cores taken from the same tree and with cores taken from different trees. these similarities increased with tree size. The climatic change affected the large trees more strongly than it did the small trees. Micro environmental changes, such as canopy gap affected the small trees more strongly. Annual ring widths were also correlated with the annual precipitations at Jumla 30 km south of the plots for a recent 20 year period. A multiple regression analysis between ring width and seasonal precipitation showed that the growth ofA. spectabilis was correlated primarily with the rain from May to August and secondarily with that from September to December in the previous year.     

Growth and resilience responses of Scots pine to extreme droughts across Europe depend on predrought growth conditions

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.     

The energy–water limitation threshold explains divergent drought responses in tree growth,needle length,and stable isotope ratios

Predicted increases in extreme droughts will likely cause major shifts in carbon sequestration and forest composition. Although growth declines during drought are widely documented, an increasing number of studies have reported both positive and negative responses to the same drought. These divergent growth patterns may reflect thresholds (i.e., nonlinear responses) promoted by changes in the dominant climatic constraints on tree growth. Here we tested whether stemwood growth exhibited linear or nonlinear responses to temperature and precipitation and whether stemwood growth thresholds co-occurred with multiple thresholds in source and sink processes that limit tree growth. We extracted 772 tree cores, 1398 needle length records, and 1075 stable isotope samples from 27 sites across whitebark pine's (Pinus albicaulis Engelm.) climatic niche in the Sierra Nevada. Our results indicated that a temperature threshold in stemwood growth occurred at 8.4°C (7.12–9.51°C; estimated using fall-spring maximum temperature). This threshold was significantly correlated with thresholds in foliar growth, as well as carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope ratios, that emerged during drought. These co-occurring thresholds reflected the transition between energy- and water-limited tree growth (i.e., the E–W limitation threshold). This transition likely mediated carbon and nutrient cycling, as well as important differences in growth-defense trade-offs and drought adaptations. Furthermore, whitebark pine growing in energy-limited regions may continue to experience elevated growth in response to climate change. The positive effect of warming, however, may be offset by growth declines in water-limited regions, threatening the long-term sustainability of the recently listed whitebark pine species in the Sierra Nevada.     

Know Your Neighbours: Drought Response of Norway Spruce,Silver Fir and Douglas Fir in Mixed Forests Depends on Species Identity and Diversity of Tree Neighbourhoods

Norway spruce is a widely cultivated species in Central Europe; however, it is highly susceptible to droughts, which are predicted to become more frequent in the future. A solution to adapt spruce forests to droughts could be the conversion to mixed-species stands containing species which are less sensitive to drought and do not increase the drought stress in spruce. Here we assessed the drought response of spruce and the presumably more drought-tolerant silver fir and Douglas fir in mixed-conifer stands. We measured tree ring widths of 270 target trees, which grew in mixed and mono-specific neighbourhoods in 18 managed stands in the Black Forest, to quantify the complementarity effects caused by species interactions on growth during the extreme drought event of 2003 and for a number of years with “normal” growth and climatic conditions. Mixed-species neighbourhoods did not significantly affect tree ring growth in normal years. However, during the drought, silver fir benefitted from mixing, while Douglas fir was more drought-stressed in the mixture. The drought response of spruce was dependent on the density and species composition of the neighbourhood, showing both positive and negative mixing effects. Mixed stands containing these tree species could improve adaptation to drought because the risks of extreme events are spread across species, and the performance of individual species is improved. Our knowledge about specific species interactions needs to be improved to manage tree mixtures more effectively with regard to the participating species and stand density.     

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.     

Exposure of trees to drought‐induced die‐off is defined by a common climatic threshold across different vegetation types

Increases in drought and temperature stress in forest and woodland ecosystems are thought to be responsible for the rise in episodic mortality events observed globally. However, key climatic drivers common to mortality events and the impacts of future extreme droughts on tree survival have not been evaluated. Here, we characterize climatic drivers associated with documented tree die‐off events across Australia using standardized climatic indices to represent the key dimensions of drought stress for a range of vegetation types. We identify a common probabilistic threshold associated with an increased risk of die‐off across all the sites that we examined. We show that observed die‐off events occur when water deficits and maximum temperatures are high and exist outside 98% of the observed range in drought intensity; this threshold was evident at all sites regardless of vegetation type and climate. The observed die‐off events also coincided with at least one heat wave (three consecutive days above the 90th percentile for maximum temperature), emphasizing a pivotal role of heat stress in amplifying tree die‐off and mortality processes. The joint drought intensity and maximum temperature distributions were modeled for each site to describe the co‐occurrence of both hot and dry conditions and evaluate future shifts in climatic thresholds associated with the die‐off events. Under a relatively dry and moderate warming scenario, the frequency of droughts capable of inducing significant tree die‐off across Australia could increase from 1 in 24 years to 1 in 15 years by 2050, accompanied by a doubling in the occurrence of associated heat waves. By defining commonalities in drought conditions capable of inducing tree die‐off, we show a strong interactive effect of water and high temperature stress and provide a consistent approach for assessing changes in the exposure of ecosystems to extreme drought events.     

Tree ring isotopic composition, radial increment and height growth reveal provenance-specific reactions of Douglas-fir towards environmental parameters

In the search of timber species being tolerant towards summer droughts, which are expected to be more frequent in future, Douglas-fir is often discussed as a potential alternative for spruce in Central Europe. To assess physiological and growth reactions of Douglas-fir provenances towards climate- and weather-related environmental conditions we took advantage of a provenance trial with three sites in south-western Germany located along an elevation gradient. We examined six different provenances of Douglas-fir from North America for oxygen (δ¹⁸O) and carbon (δ¹³C) stable isotope composition in tree rings as well as for radial increment for a 7 year period and long-term height growth. Our results show that different Douglas-fir provenances clearly vary in their drought sensitivity at the driest and warmest site in the valley as shown by the radial growth decline in the extreme dry and hot year 2003. The growth decline in the provenances Pamelia Creek, Cameron Lake, Duncan Paldi and Conrad Creek could be clearly attributed to a reduction in stomatal conductance as assessed by the relations between δ¹⁸O and δ¹³C in the tree rings. These responses were not related to the long-term average climate at the places of origin of the provenances and the provenances with the lowest long-term (height) growth potential were the ones least affected in radial increment by the extreme drought of 2003. When selecting suitable Douglas-fir provenances, which are adapted to the climatic conditions projected for the future we thus might need to take into account the trade-off between the adaptation to extreme drought periods and the long-term growth performance. Site-specific evaluations of the probability of extreme drought events are thus needed to select the appropriate provenances.     

Resistance of European tree species to drought stress in mixed versus pure forests: evidence of stress release by inter‐specific facilitation

While previous studies focused on tree growth in pure stands, we reveal that tree resistance and resilience to drought stress can be modified distinctly through species mixing. Our study is based on tree ring measurement on cores from increment boring of 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in South Germany, with half sampled in pure, respectively, mixed stands. Indices for resistance, recovery and resilience were applied for quantifying the tree growth reaction on the episodic drought stress in 1976 and 2003. The following general reaction patterns were found. (i) In pure stands, spruce has the lowest resistance, but the quickest recovery; oak and beech were more resistant, but recover was much slower and they are less resilient. (ii) In mixture, spruce and oak perform as in pure stands, but beech was significantly more resistant and resilient than in monoculture. (iii) Especially when mixed with oak, beech is facilitated. We hypothesise that the revealed water stress release of beech emerges in mixture because of the asynchronous stress reaction pattern of beech and oak and a facilitation of beech by hydraulic lift of water by oak. This facilitation of beech in mixture with oak means a contribution to the frequently reported overyield of beech in mixed versus pure stands. We discuss the far‐reaching implications that these differences in stress response under intra‐ and inter‐specific environments have for forest ecosystem dynamics and management under climate change.     

Long‐term successional forest dynamics: species and community responses to climatic variability

Question: Are trees sensitive to climatic variability, and do tree species differ in their responses to climatic variability? Does sensitivity of forest communities to climatic variability depend on stand composition? Location: Mixed young forest at Walker Branch Watershed near Oak Ridge, East Tennessee, USA. Methods: Using a long‐term dataset (1967–2006), we analyzed temporal forest dynamics at the tree and species level, and community dynamics for forest stands that differed in initial species composition (i.e., chestnut oak, oak–hickory, pine, and yellow poplar stands). Using summer drought and growing season temperature as defined climate drivers, we evaluated relationships between forest dynamics and climate across levels of organization. Results: Over the four‐decade study period, forest communities underwent successional change and substantially increased in biomass. Variation in summer drought and growing season temperature contributed to temporal biomass dynamics for some tree species, but not for others. Stand‐level responses to climatic variability were related to the responses of component species, except in pine stands. Pinus echinata, the dominant species in pine stands, decreased over time due to periodic outbreaks of pine bark beetle (Dendroctonus frontalis). These outbreaks at Walker Branch could not be directly related to climatic conditions. Conclusions: The results indicate that sensitivity of developing forests to climatic variability is stand type‐dependent, and hence is a function of species composition. However, in the long term, direct effects of climatic variability on forest dynamics may be small relative to autogenic successional processes or climate‐related insect outbreaks. Empirical studies testing for interactions between forest succession and climatic variability are needed.