干旱和林分因子对树木死亡的影响——以美国德克萨斯州东部国家森林为例

为探讨不同时间尺度、气候因子及林分因子对森林中树木死亡的影响,本研究以美国德克萨斯州东部的4个国家森林中264个重复调查的森林样地为对象,使用近20年来美国森林清查4个周期的数据,估算其在清查周期和年度水平上的树木死亡率变化,并使用广义线性混合效应模型来分析气候因子(干旱强度、干旱持续时间、年均温和年降水量)、树木大小(胸径)和林分因子(树木胸高断面积、林分密度和林分年龄)对树木存活的影响。结果表明: 在重度干旱当年和重度干旱的清查周期中,森林的树木死亡率分别增加了151%和123%,天气干扰(干旱和飓风)和植物之间的竞争是其主要的影响因素;干旱强度(标准化降水蒸散发指数,SPEI)和干旱持续时间对树木的存活具有显著的负效应,年降水量对树木的存活具有显著的正效应;树木胸高面积对树木存活具有显著的负效应,树木大小、林分年龄和林分密度对树木存活均具有显著的正效应,但是大树比小树更容易受到天气影响而死亡;在重度干旱的清查周期中,松树种组的树木死亡率(2.1%)比阔叶树木种组(3.9%)低,天然林的树木死亡率(3.0%)高于人工林(1.9%)。在分析树木死亡率时,需同时考虑个体树木大小、林分因子与气候因子的相对重要性。     

Drought and the interannual variability of stem growth in an aseasonal,everwet forest

Linking drought to the timing of physiological processes governing tree growth remains one limitation in forecasting climate change effects on tropical trees. Using dendrometers, we measured fine‐scale growth for 96 trees of 25 species from 2013 to 2016 in an everwet forest in Puerto Rico. Rainfall over this time span varied, including an unusual, severe El Niño drought in 2015. We assessed how growing season onset, median day, conclusion, and length varied with absolute growth rate and tree size over time. Stem growth was seasonal, beginning in February, peaking in July, and ending in November. Species growth rates varied between 0 and 8 mm/year and correlated weakly with specific leaf area, leaf phosphorus, and leaf nitrogen, and to a lesser degree with wood specific gravity and plant height. Drought and tree growth were decoupled, and drought lengthened and increased variation in growing season length. During the 2015 drought, many trees terminated growth early but did not necessarily grow less. In the year following drought, trees grew more over a shorter growing season, with many smaller trees showing a post‐drought increase in growth. We attribute the increased growth of smaller trees to release from light limitation as the canopy thinned because of the drought, and less inferred hydraulic stress than larger trees during drought. Soil type accounted for interannual and interspecific differences, with the finest Zarzal clays reducing tree growth. We conclude that drought affects the phenological timing of tree growth and favors the post‐drought growth of smaller, sub‐canopy trees in this everwet forest. Abstract in Spanish is available with online material.     

4种阔叶幼苗对PEG模拟干旱的生理响应

研究了PEG模拟干旱胁迫环境下的火力楠(Michelia macclurel)、尾叶桉(Eucalyptus urophylla)、枫香(Liquidambar formosana)、荷木(Schima superba)幼苗的生理变化。结果表明,干旱胁迫下,4种幼苗叶片的相对含水量小于对照,其中,尾叶桉和枫香下降明显;不同干旱胁迫条件下,4种树种幼苗叶片的相对电导率均显著大于对照,其中尾叶桉和枫香上升幅度大;干旱胁迫下的火力楠和荷木幼苗叶片的脯氨酸含量呈现波动,尾叶桉和枫香幼苗则显著大于对照;不同干旱胁迫时间下的幼苗叶片的叶绿素含量小幅波动;4个树种幼苗的过氧岐化酶(SOD)活性随胁迫时间增加而呈现先升后降的趋势,其中火力楠和荷木的幼苗的SOD活性持续维持在较高水平;荷木叶片的丙二醛(MDA)含量先升后降,最后和对照水平相近,其余幼苗的MDA含量均大于对照;干旱胁迫下4种幼苗叶片的可溶性糖含量增加幅度较大。主成分分析表明,4种幼苗的抗旱能力排序为荷木>火力楠>尾叶桉>枫香。     

Drought-induced tree growth decline in the desert margins of Northwestern China

Water deficiency is the primary limiting factor for tree growth in arid and semi-arid areas. Droughts associated with rising temperatures have increased in severity and frequency globally over the past few decades, making the trees in the drought-prone sites first be affected by water shortages. However, our understanding of tree growth status in these areas, and of their response to drought, is currently insufficient; especially in the context of global warming. Here, we studied 94 Chinese pine (Pinus tabulaeformis) and 86 spruce (Picea crassifolia) trees from different altitudes [2,100–2400 m above sea level (a.s.l.)] distributed at the desert margins of Northwestern China to explore tree growth and drought response from multiple perspectives using dendroecological approaches. Significant growth decline, across all tree species and altitudes, was detected in response to an interdecadal trend towards a drier climate. Moreover, the extent of tree growth decline, the proportion of affected trees, and the degree of moisture dependence have all tended to increase in each sample site, most likely due to enhanced drought severity and duration in recent decades. The more sensitive and susceptible trees were found at lower elevations (drier sites) and may signify a higher vulnerability to heating-induced drought stress. Tree resistance to drought showed strong negative correlation with drought severity across all sample sites. However, the connection between post-drought tree resilience and drought intensity is weak, perhaps because the samples were all collected from living trees, while those that had died were not sampled. The priority for future work should be to combine surviving and dead trees simultaneously, thus achieving a more representative view of tree resilience to drought; this will improve our knowledge of forest dynamics and even ecosystem succession in these vulnerable and sensitive environments.     

Drought as an Inciting Mortality Factor in Scots Pine Stands of the Valais, Switzerland

During the 20th century, high mortality rates of Scots pine (Pinus silvestris L.) have been observed over large areas in the Rhône valley (Valais, Switzerland) and in other dry valleys of the European Alps. In this study, we evaluated drought as a possible inciting factor of Scots pine decline in the Valais. Averaged tree-ring widths, standardized tree-ring series, and estimated annual mortality risks were related to a drought index. Correlations between drought indices and standardized tree-ring series from 11 sites showed a moderate association. Several drought years and drought periods could be detected since 1864 that coincided with decreased growth. Although single, extreme drought years had generally a short-term, reversible effect on tree growth, multi-year drought initiated prolonged growth decreases that increased a tree’s long-term risk of death. Tree death occurred generally several years or even decades after the drought. In conclusion, drought has a limiting effect on tree growth and acts as a bottleneck event in triggering Scots pine decline in the Valais.     

Tree carbon allocation explains forest drought‐kill and recovery patterns

The mechanisms governing tree drought mortality and recovery remain a subject of inquiry and active debate given their role in the terrestrial carbon cycle and their concomitant impact on climate change. Counter‐intuitively, many trees do not die during the drought itself. Indeed, observations globally have documented that trees often grow for several years after drought before mortality. A combination of meta‐analysis and tree physiological models demonstrate that optimal carbon allocation after drought explains observed patterns of delayed tree mortality and provides a predictive recovery framework. Specifically, post‐drought, trees attempt to repair water transport tissue and achieve positive carbon balance through regrowing drought‐damaged xylem. Furthermore, the number of years of xylem regrowth required to recover function increases with tree size, explaining why drought mortality increases with size. These results indicate that tree resilience to drought‐kill may increase in the future, provided that CO₂ fertilisation facilitates more rapid xylem regrowth.     

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.     

Testing early warning metrics for drought‐induced tree physiological stress and mortality

Climate change‐driven drought stress has triggered numerous large‐scale tree mortality events in recent decades. Advances in mechanistic understanding and prediction are greatly limited by an inability to detect in situ where trees are likely to die in order to take timely measurements and actions. Thus, algorithms of early warning and detection of drought‐induced tree stress and mortality could have major scientific and societal benefits. Here, we leverage two consecutive droughts in the southwestern United States to develop and test a set of early warning metrics. Using Landsat satellite data, we constructed early warning metrics from the first drought event. We then tested these metrics' ability to predict spatial patterns in tree physiological stress and mortality from the second drought. To test the broader applicability of these metrics, we also examined a separate drought in the Amazon rainforest. The early warning metrics successfully explained subsequent tree mortality in the second drought in the southwestern US, as well as mortality in the independent drought in tropical forests. The metrics also strongly correlated with spatial patterns in tree hydraulic stress underlying mortality, which provides a strong link between tree physiological stress and remote sensing during the severe drought and indicates that the loss of hydraulic function during drought likely mediated subsequent mortality. Thus, early warning metrics provide a critical foundation for elucidating the physiological mechanisms underpinning tree mortality in mature forests and guiding management responses to these climate‐induced disturbances.     

Different responses of multispecies tree ring growth to various drought indices across Europe

Increasing frequency and intensity of drought extremes associated with global change are a key challenge for forest ecosystems. Consequently, the quantification of drought effects on tree growth as a measure of vitality is of highest concern from the perspectives of both science and management. To date, a multitude of drought indices have been used to accompany or replace primary climatic variables in the analysis of drought-related growth responses. However, it remains unclear how individual drought metrics compare to each other in terms of their ability to capture drought signals in tree growth.In our study, we employ a European multispecies tree ring network at the continental scale and a set of four commonly used drought indices (De Martonne Aridity Index, self-calibrating Palmer Drought Severity Index, Standardized Precipitation Index and Standardized Precipitation Evapotranspiration Index, the latter two on varying temporal scales) to derive species-specific growth responses to drought conditions. For nine common European tree species, we demonstrate spatio-temporal matches and mismatches of tree growth with drought indices subject to species, elevation and bioclimatic zone. Forests located in the temperate and Mediterranean climate were drought sensitive and tended to respond to short- and intermediate-term drought (<1 year). In continental climates, forests were comparably more drought resistant and responded to long-term drought. For the same species, stands were less drought sensitive at higher elevations compared to lower elevations. We provide detailed information on the month-wise performance of the four drought indices in different climate zones allowing users the selection of the most appropriate index according to their objective criteria. Our results show that species-specific differences in responses to multiple stressors result in complex, yet coherent patterns of tree growth.     

特大干旱对树木死亡的影响——以美国德克萨斯州东部森林为例

美国德克萨斯州在2011年经历了史上最严重的干旱,这一事件造成约3亿多株树木死亡。在大时空尺度上(面积约9×10~6 hm~2,时间跨度近20年),基于近1800个森林样地,4次周期性调查中的约209663株树木,使用主成份分析(PCA)和广义线性混合效应模型(GLM)回归,对树木死亡的时空差异及其干旱强度与长度对树木死亡造成的中长期复杂影响进行了研究。采用树木密度、树木基面积、林地年龄、样地调查时间间隔作为树木间的竞争指标,分析了造成大旱前后周期水平和年度水平上的样地树木死亡差异的原因。综合分析了不同地理区域、树木种组、胸径大小和林地起源的4个划分标准下树木死亡对死亡率的相对贡献。结果表明：松属树木的死亡率最低(7.92%);高度低、胸径小的树木的死亡率较大,分别为29.79%和26.00%。人工林的树木死亡率(10.26%)低于天然林(13.47%);西海湾平原生态区树木的死亡率在干旱后达到最大(22.27%);西南区的树木死亡率在干旱后也达到最大(13.78%);海拔和纬度对树木死亡率影响不明显。德州东部森林整体死亡格局形成原因较为复杂,各地理区域、林地起源、树木大小和不同树种,...     

Drought‐related tree death of savanna eucalypts: Species susceptibility,soil conditions and root architecture

Questions: For eucalypt savanna in northeast Australia subject to multi‐year rainfall deficits this paper asks whether (1) dominant tree species (Ironbarks, Boxes) are more drought susceptible than the sub‐dominant Bloodwoods; (2) whether soil moisture is beyond wilting point in surface soil layers but available at depth; (3) soil conditions (moisture availability and texture) are related to tree death during drought; (4) the root systems of the Boxes and Ironbarks are shallower than the Bloodwoods; and the survivors of drought within species have deeper root systems than those that died. Location: Central Queensland, Australia. Methods: Patterns of tree death between eucalypt species were compared from field data collected after drought. Soil conditions during drought were described and compared with patterns of tree death for the Ironbark Eucalyptus melanophloia. The basal area and orientation of coarse roots were measured on upturned trees after broad‐scale tree clearing, and compared between species, and between live and dead trees with tree size as a covariate. Results: Drought‐induced tree death was higher for dominant Ironbark‐Box than for sub‐dominant Bloodwoods. During a moderate to severe drought in 2004, 41% of 100 cm deep subsoils had soil matric potential less than‐5600 kPa. The drought hardy Bloodwoods had a greater root basal area and particularly so for vertical roots compared to the drought sensitive Ironbark‐Box. Within species there was no significant difference in root basal area characteristics between trees that were recently killed by drought and those that remained relatively healthy. Surface soil moisture availability was lower where tree densities were high, and tree death increased as surface soil moisture became less available. Tree death was also greater as the clay content of sub‐soils increased. Discussion: The study suggests species with roots confined to upper soil layers will suffer severe water stress. The results strongly indicate that root architecture, and the way it facilitates water use during drought, is important for the relative dominance of the tree species. Patchiness in drought‐induced tree death seems to be at least partially a product of heterogeneity in sub‐soil conditions and competition for soil moisture.     

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.     

The worst drought in 50 years in a South African savannah: Limited impact on vegetation

South Africa experienced a severe multiyear drought from 2014 to 2016. Here, we explore the response of a South African savannah ecosystem to this drought focusing on tree and grass dynamics. We used open long‐term monitoring plots established in 2000 and distributed across broad rainfall gradients in the Hluhluwe‐iMfolozi Park in KwaZulu‐Natal. Analysis showed negligible tree mortality due to drought in 2016 (0.03%) and 2017 (0.49%). However, there was an apparent increase in the tree population especially among the small size classes (0.1–0.3 m) in 2016 and 2017 relative to the predrought 2012 census. Drought effects may be confounded with changes in browser populations. The impala population declined in a mesic savannah (Hluhluwe), as measured with dung counts; however, this decline started before the drought. Impala decline in a semi‐arid savannah (iMfolozi) was more coincident with drought. Grass biomass and cover decreased in 2016 compared to predrought 2012 but showed rapid recovery once rains began. In iMfolozi grass species, composition improved from a grazing perspective as a result of the recovery of the decreaser species, an increase in palatable species and a marked decline of unpalatable species. The minor changes in woody plants and grasses from before the drought until rain resumed and the rapid recovery of grasses suggest that this South African savannah ecosystem was resilient to severe drought.     

Can tree species richness attenuate the effect of drought on organic matter decomposition and stabilization in young plantation forests?

Changes in precipitation due to climate change are likely to influence soil organic matter (SOM) decomposition and stabilization. In forests, increased tree species diversity could modulate the effects of drought on SOM decomposition and stabilization. We addressed this issue by a decomposition study under simulated drought (through precipitation reduction at Zedelgem, Belgium) and natural drought (ORPHEE, southern France) in young experimental plantations (tree species richness 1 to 5). In Zedelgem, the study focused on tree species richness around oak and beech trees. Two tea bag indices (TBI) – decomposition rate (k) and stabilization factor (S) – were calculated by measuring the decay of green and rooibos tea in soils. Overall, TBI's were higher in Zedelgem than at ORPHEE. In Zedelgem, k increased with tree species richness under drought around oak, indicating that tree species richness modulated the effects of drought on decomposition. Under beech, k decreased with drought while no effect of tree species richness was detected. S increased with drought under both oak and beech, without any effect of tree species richness. In ORPHEE, we did not detect any tree species richness effect on both TBIs. S decreased significantly, while k was marginally reduced under drought. The higher S under drought in Zedelgem and under control in OPRHEE suggests that the carbon sequestration potential under climate change would be dependent on the environmental context. Further, in young plantations, high species richness may modulate the drought effect on SOM decomposition, but not on stabilization.     

El Niño droughts and their effects on tree species composition and diversity in tropical rain forests

In this study I investigated the effects of the extreme, 1997/98 El Niño related drought on tree mortality and understorey light conditions of logged and unlogged tropical rain forest in the Indonesian province of East Kalimantan (Borneo). My objectives were to test (1) whether drought had a significant effect on tree mortality and understorey light conditions, (2) whether this effect was greater in logged than in undisturbed forest, (3) if the expected change in tree mortality and light conditions had an effect on Macaranga pioneer seedling and sapling densities, and (4) which (a)biotic factors influenced tree mortality during the drought. The 1997/1998 drought led to an additional tree mortality of 11.2, 18.1, and 22.7% in undisturbed, old logged and recently logged forest, respectively. Mortality was highest in logged forests, due to extremely high mortality of pioneer Macaranga trees (65.4%). Canopy openness was significantly higher during the drought than during the non-drought year (6.0, 8.6 and 10.4 vs 3.7, 3.8 and 3.7 in undisturbed, old logged and recently logged forest, respectively) and was positively correlated with the number of dead standing trees. The increase in light in the understorey was accompanied by a 30 to 300-fold increase in pioneer Macaranga seedling densities. Factors affecting tree mortality during drought were (1) tree species successional status, (2) tree size, and (3) tree location with respect to soil moisture. Tree density and basal area per surface unit had no influence on tree mortality during drought. The results of this study show that extreme droughts, such as those associated with El Niño events, can affect the tree species composition and diversity of tropical forests in two ways: (1) by disproportionate mortality of certain tree species groups and tree size classes, and (2) by changing the light environment in the forest understorey, thereby affecting the recruitment and growth conditions of small and immature trees.     

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。     

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.     

土壤干旱对黄土高原3个常见树种幼苗水分代谢及生长的影响

应用盆栽试验,在人工控制土壤水分条件下对黄土高原3个常见树种丁香(Syringa oblata)、杠柳(Perip-loca sepium)和连翘(Forsythia suspensa)幼苗的生长及水分生理代谢进行了研究.结果表明,随干旱胁迫程度加剧,各树种耗水量明显减少;不同树种单株耗水量差异明显,表现为:连翘>杠柳>丁香.3树种新生枝条生长和叶面积扩展速率明显受土壤含水量影响,均表现为适宜水分>中度干旱>严重干旱,且在同一胁迫水平下,连翘>杠柳>丁香.随干旱胁迫程度的加剧和干旱时间的延长,丁香、杠柳和连翘叶片的含水量、游离脯氨酸以及叶绿素含量均有不同程度的变化,连翘和杠柳的叶片含水量在3种水分条件下均明显高于丁香,杠柳叶片游离脯氨酸含量明显高于丁香和连翘,连翘体内脯氨酸含量最低,丁香和连翘的叶绿素a/b值随土壤含水量的减少逐渐降低,杠柳则表现出相反趋势.不同树种对土壤干旱和高温的响应机制不同,但它们都具有较强的抗旱能力,适应黄土高原干旱的自然条件.     

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.     

Long-Term Tree Cover Dynamics in a Pinyon-Juniper Woodland: Climate-Change-Type Drought Resets Successional Clock

Woody vegetation has expanded in coverage over the past century in many places globally, exemplified by pinyon-juniper changes in the Southwestern United States. Extreme drought is one of the few non-management drivers besides fire that might reverse such cover changes, but this has not been well documented. Here, we assess 68 years of tree cover dynamics across an elevation gradient of a pinyon-juniper woodland using aerial photographs (1936 and 1959) and QuickBird imagery (2004). Canopy cover increased 32% from 1936 to the onset of a major drought (2002). The largest relative increase in canopy cover occurred from 1936 to 1959 at the higher elevations, but these gains were eliminated by fires occurring from 1959 to 2002, during which time lower elevations with low canopy cover exhibited the greatest relative increases. The 2002–2004 drought reduced canopy cover by 55%, which eliminated gains in cover that occurred since 1936. Relative tree cover loss was highest at low elevations with low tree cover, but absolute tree cover loss was greater in areas of high tree cover, which increased with elevation. The loss of more than half of the canopy cover during a 2-year drought period was much greater than losses due to fire or possible increases due to historic land use (for example, grazing). These results suggest that regional-scale climatic influences may be more important than land use legacies in controlling tree cover of these and perhaps other semiarid woodlands over longer time scales—notable given that similar episodes of tree mortality are projected in coming decades with climate change.