Long‐term climate and competition explain forest mortality patterns under extreme drought

Rising temperatures are amplifying drought‐induced stress and mortality in forests globally. It remains uncertain, however, whether tree mortality across drought‐stricken landscapes will be concentrated in particular climatic and competitive environments. We investigated the effects of long‐term average climate [i.e. 35‐year mean annual climatic water deficit (CWD)] and competition (i.e. tree basal area) on tree mortality patterns, using extensive aerial mortality surveys conducted throughout the forests of California during a 4‐year statewide extreme drought lasting from 2012 to 2015. During this period, tree mortality increased by an order of magnitude, typically from tens to hundreds of dead trees per km², rising dramatically during the fourth year of drought. Mortality rates increased independently with average CWD and with basal area, and they increased disproportionately in areas that were both dry and dense. These results can assist forest managers and policy‐makers in identifying the most drought‐vulnerable forests across broad geographic areas.     

Chronic water stress reduces tree growth and the carbon sink of deciduous hardwood forests

Predicted decreases in water availability across the temperate forest biome have the potential to offset gains in carbon (C) uptake from phenology trends, rising atmospheric CO₂, and nitrogen deposition. While it is well established that severe droughts reduce the C sink of forests by inducing tree mortality, the impacts of mild but chronic water stress on forest phenology and physiology are largely unknown. We quantified the C consequences of chronic water stress using a 13‐year record of tree growth (n = 200 trees), soil moisture, and ecosystem C balance at the Morgan–Monroe State Forest (MMSF) in Indiana, and a regional 11‐year record of tree growth (n > 300 000 trees) and water availability for the 20 most dominant deciduous broadleaf tree species across the eastern and midwestern USA. We show that despite ~26 more days of C assimilation by trees at the MMSF, increasing water stress decreased the number of days of wood production by ~42 days over the same period, reducing the annual accrual of C in woody biomass by 41%. Across the deciduous forest region, water stress induced similar declines in tree growth, particularly for water‐demanding ‘mesophytic’ tree species. Given the current replacement of water‐stress adapted ‘xerophytic’ tree species by mesophytic tree species, we estimate that chronic water stress has the potential to decrease the C sink of deciduous forests by up to 17% (0.04 Pg C yr^?1) in the coming decades. This reduction in the C sink due to mesophication and chronic water stress is equivalent to an additional 1–3 days of global C emissions from fossil fuel burning each year. Collectively, our results indicate that regional declines in water availability may offset the growth‐enhancing effects of other global changes and reduce the extent to which forests ameliorate climate warming.     

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,...     

Growth, carbon-isotope discrimination, and drought-associated mortality across a Pinus ponderosa elevational transect

Drought‐ and insect‐associated tree mortality at low‐elevation ecotones is a widespread phenomenon but the underlying mechanisms are uncertain. Enhanced growth sensitivity to climate is widely observed among trees that die, indicating that a predisposing physiological mechanism(s) underlies tree mortality. We tested three, linked hypotheses regarding mortality using a ponderosa pine (Pinus ponderosa) elevation transect that experienced low‐elevation mortality following prolonged drought. The hypotheses were: (1) mortality was associated with greater growth sensitivity to climate, (2) mortality was associated with greater sensitivity of gas exchange to climate, and (3) growth and gas exchange were correlated. Support for all three hypotheses would indicate that mortality results at least in part from gas exchange constraints. We assessed growth using basal area increment normalized by tree basal area [basal area increment (BAI)/basal area (BA)] to account for differences in tree size. Whole‐crown gas exchange was indexed via estimates of the CO₂ partial pressure difference between leaf and atmosphere (p_a?p_c) derived from tree ring carbon isotope ratios (δ¹³C), corrected for temporal trends in atmospheric CO₂ and δ¹³C and elevation trends in pressure. Trees that survived the drought exhibited strong correlations among and between BAI, BAI/BA, p_a?p_c, and climate. In contrast, trees that died exhibited greater growth sensitivity to climate than trees that survived, no sensitivity of p_a?p_c to climate, and a steep relationship between p_a?p_c and BAI/BA. The p_a?p_c results are consistent with predictions from a theoretical hydraulic model, suggesting trees that died had a limited buffer between mean water availability during their lifespan and water availability during drought – i.e., chronic water stress. It appears that chronic water stress predisposed low‐elevation trees to mortality during drought via constrained gas exchange. Continued intensification of drought in mid‐latitude regions may drive increased mortality and ecotone shifts in temperate forests and woodlands.     

Cocoa agroforestry is less resilient to sub‐optimal and extreme climate than cocoa in full sun

Cocoa agroforestry is perceived as potential adaptation strategy to sub‐optimal or adverse environmental conditions such as drought. We tested this strategy over wet, dry and extremely dry periods comparing cocoa in full sun with agroforestry systems: shaded by (i) a leguminous tree species, Albizia ferruginea and (ii) Antiaris toxicaria, the most common shade tree species in the region. We monitored micro‐climate, sap flux density, throughfall, and soil water content from November 2014 to March 2016 at the forest‐savannah transition zone of Ghana with climate and drought events during the study period serving as proxy for projected future climatic conditions in marginal cocoa cultivation areas of West Africa. Combined transpiration of cocoa and shade trees was significantly higher than cocoa in full sun during wet and dry periods. During wet period, transpiration rate of cocoa plants shaded by A. ferruginea was significantly lower than cocoa under A. toxicaria and full sun. During the extreme drought of 2015/16, all cocoa plants under A. ferruginea died. Cocoa plants under A. toxicaria suffered 77% mortality and massive stress with significantly reduced sap flux density of 115 g cm^?2 day^?1, whereas cocoa in full sun maintained higher sap flux density of 170 g cm^?2 day^?1. Moreover, cocoa sap flux recovery after the extreme drought was significantly higher in full sun (163 g cm^?2 day^?1) than under A. toxicaria (37 g cm^?2 day^?1). Soil water content in full sun was higher than in shaded systems suggesting that cocoa mortality in the shaded systems was linked to strong competition for soil water. The present results have major implications for cocoa cultivation under climate change. Promoting shade cocoa agroforestry as drought resilient system especially under climate change needs to be carefully reconsidered as shade tree species such as the recommended leguminous A. ferruginea constitute major risk to cocoa functioning under extended severe drought.     

A retrospective,dual‐isotope approach reveals individual predispositions to winter‐drought induced tree dieback in the southernmost distribution limit of Scots pine

Winter‐drought induced forest diebacks in the low‐latitude margins of species' distribution ranges can provide new insights into the mechanisms (carbon starvation, hydraulic failure) underlying contrasting tree reactions. We analysed a winter‐drought induced dieback at the Scots pine's southern edge through a dual‐isotope approach (Δ¹³C and δ¹⁸O in tree‐ring cellulose). We hypothesized that a differential long‐term performance, mediated by the interaction between CO₂ and climate, determined the fates of individuals during dieback. Declining trees showed a stronger coupling between climate, growth and intrinsic water‐use efficiency (WUEi) than non‐declining individuals that was noticeable for 25 years prior to dieback. The rising stomatal control of water losses with time in declining trees, indicated by negative Δ¹³C‐δ¹⁸O relationships, was likely associated with their native aptitude to grow more and take up more water (suggested by larger tracheid lumen widths) than non‐declining trees and, therefore, to exhibit a greater cavitation risk. Freeze‐thaw episodes occurring in winter 2001 unveiled such physiological differences by triggering dieback in those trees more vulnerable to hydraulic failure. Thus, WUEi tightly modulated growth responses to long‐term warming in declining trees, indicating that co‐occurring individuals were differentially predisposed to winter‐drought mortality. These different performances were unconnected to the depletion of stored carbohydrates.     

Climate change‐associated tree mortality increases without decreasing water availability

Temporal increases of tree mortality have been observed in regions where global warming has decreased long‐term water availability and/or induced droughts. However, temporal decreases in water availability are not a global phenomenon. Understanding how water deficit‐free forests respond to the recent effects of climate change is paramount towards a full appreciation of the impacts of climate change on global forests. Here, we reveal temporally increasing tree mortality across all study species over the last three decades in the central boreal forests of Canada, where long‐term water availability has increased without apparent climate change‐associated drought. In addition, we find that the effects of conspecific tree‐to‐tree competition have intensified temporally as a mechanism for the increased mortality of shade‐intolerant tree species. Our results suggest that the consequences of climate change on tree mortality are more profound than previously thought.     

Temperature‐induced water stress in high‐latitude forests in response to natural and anthropogenic warming

The Arctic is particularly sensitive to climate change, but the independent effects of increasing atmospheric CO₂ concentration (pCO₂) and temperature on high‐latitude forests are poorly understood. Here, we present a new, annually resolved record of stable carbon isotope (δ¹³C) data determined from Larix cajanderi tree cores collected from far northeastern Siberia in order to investigate the physiological response of these trees to regional warming. The tree‐ring record, which extends from 1912 through 1961 (50 years), targets early twentieth‐century warming (ETCW), a natural warming event in the 1920s to 1940s that was limited to Northern hemisphere high latitudes. Our data show that net carbon isotope fractionation (Δ¹³C), decreased by 1.7‰ across the ETCW, which is consistent with increased water stress in response to climate warming and dryer soils. To investigate whether this signal is present across the northern boreal forest, we compiled published carbon isotope data from 14 high‐latitude sites within Europe, Asia, and North America. The resulting dataset covered the entire twentieth century and spanned both natural ETCW and anthropogenic Late Twentieth‐Century Warming (~0.7 °C per decade). After correcting for a ~1‰ increase in Δ¹³C in response to twentieth century pCO₂ rise, a significant negative relationship (r = ?0.53, P < 0.0001) between the average, annual Δ¹³C values and regional annual temperature anomalies is observed, suggesting a strong control of temperature on the Δ¹³C value of trees growing at high latitudes. We calculate a 17% increase in intrinsic water‐use efficiency within these forests across the twentieth century, of which approximately half is attributed to a decrease in stomatal conductance in order to conserve water in response to drying conditions, with the other half being attributed to increasing pCO₂. We conclude that annual tree‐ring records from northern high‐latitude forests record the effects of climate warming and pCO₂ rise across the twentieth century.     

Transformation of riparian forest to woodland in Mapungubwe National Park,South Africa,between 1990 and 2007

Canopy tree survival and compositional change of the Greefswald forest on the Limpopo River, South Africa, were monitored between 1990 and 2007 in response to a severe drought (cessation of flow in 1991/2), water abstraction commencing in 1991, a mega‐flood in 2000 and increasing impact of elephants since 2000. Aerial photographs confirmed that forest area had not decreased during the 35 years prior to the study. In total, 25% of 428 canopy trees tagged in 1990 had died by 2005. Tree density was reduced from 22.8 to 16.3 trees per hectare. Forest was thus transformed to woodland. Mortality was attributed mainly to drought stress (47%), drought in combination with creeper infestation (30%) and the flood (21%). Of the nine main canopy species, mortality was highest for Acacia xanthophloea (56%) and Faidherbia albida (37%) mainly because of drought‐related stress, and Ficus sycamorus (25%) mainly because of the flood. Water extraction increased drought‐related mortality in the area of extraction by 45%. Creepers rendered microphyllous but not broad‐leafed species more vulnerable to drought‐induced mortality. Elephants were responsible for a further 3% mortality between 2005 and 2007. Composition has shifted towards ‘drought‐tolerant’ species not selected by elephants, namely Philenoptera violacea, Xanthocercis zambesiaca and Schotia brachypetala. Initial concern about water abstraction was eclipsed by a rapid, unpredictable concatenation of a series of rare events that transformed forest to woodland in less than 15 years.     

Prolonged experimental drought reduces plant hydraulic conductance and transpiration and increases mortality in a piñon–juniper woodland

Plant hydraulic conductance (k_s) is a critical control on whole‐plant water use and carbon uptake and, during drought, influences whether plants survive or die. To assess long‐term physiological and hydraulic responses of mature trees to water availability, we manipulated ecosystem‐scale water availability from 2007 to 2013 in a piñon pine (Pinus edulis) and juniper (Juniperus monosperma) woodland. We examined the relationship between k_s and subsequent mortality using more than 5 years of physiological observations, and the subsequent impact of reduced hydraulic function and mortality on total woody canopy transpiration (E_C) and conductance (G_C). For both species, we observed significant reductions in plant transpiration (E) and k_s under experimentally imposed drought. Conversely, supplemental water additions increased E and k_s in both species. Interestingly, both species exhibited similar declines in k_s under the imposed drought conditions, despite their differing stomatal responses and mortality patterns during drought. Reduced whole‐plant k_s also reduced carbon assimilation in both species, as leaf‐level stomatal conductance (g_s) and net photosynthesis (A_n) declined strongly with decreasing k_s. Finally, we observed that chronically low whole‐plant k_s was associated with greater canopy dieback and mortality for both piñon and juniper and that subsequent reductions in woody canopy biomass due to mortality had a significant impact on both daily and annual canopy E_C and G_C. Our data indicate that significant reductions in k_s precede drought‐related tree mortality events in this system, and the consequence is a significant reduction in canopy gas exchange and carbon fixation. Our results suggest that reductions in productivity and woody plant cover in piñon–juniper woodlands can be expected due to reduced plant hydraulic conductance and increased mortality of both piñon pine and juniper under anticipated future conditions of more frequent and persistent regional drought in the southwestern United States.     

Rapid warming accelerates tree growth decline in semi‐arid forests of Inner Asia

Forests around the world are subject to risk of high rates of tree growth decline and increased tree mortality from combinations of climate warming and drought, notably in semi‐arid settings. Here, we assess how climate warming has affected tree growth in one of the world's most extensive zones of semi‐arid forests, in Inner Asia, a region where lack of data limits our understanding of how climate change may impact forests. We show that pervasive tree growth declines since 1994 in Inner Asia have been confined to semi‐arid forests, where growing season water stress has been rising due to warming‐induced increases in atmospheric moisture demand. A causal link between increasing drought and declining growth at semi‐arid sites is corroborated by correlation analyses comparing annual climate data to records of tree‐ring widths. These ring‐width records tend to be substantially more sensitive to drought variability at semi‐arid sites than at semi‐humid sites. Fire occurrence and insect/pathogen attacks have increased in tandem with the most recent (2007–2009) documented episode of tree mortality. If warming in Inner Asia continues, further increases in forest stress and tree mortality could be expected, potentially driving the eventual regional loss of current semi‐arid forests.     

UPLC‐HRMS‐based untargeted metabolic profiling reveals changes in chickpea (Cicer arietinum) metabolome following long‐term drought stress

Genetic improvement for drought tolerance in chickpea requires a solid understanding of biochemical processes involved with different physiological mechanisms. The objective of this study is to demonstrate genetic variations in altered metabolic levels in chickpea varieties (tolerant and sensitive) grown under contrasting water regimes through ultrahigh‐performance liquid chromatography/high‐resolution mass spectrometry‐based untargeted metabolomic profiling. Chickpea plants were exposed to drought stress at the 3‐leaf stage for 25 days, and the leaves were harvested at 14 and 25 days after the imposition of drought stress. Stress produced significant reduction in chlorophyll content, F_v/F_m, relative water content, and shoot and root dry weight. Twenty known metabolites were identified as most important by 2 different methods including significant analysis of metabolites and partial least squares discriminant analysis. The most pronounced increase in accumulation due to drought stress was demonstrated for allantoin, l ‐proline, l ‐arginine, l ‐histidine, l ‐isoleucine, and tryptophan. Metabolites that showed a decreased level of accumulation under drought conditions were choline, phenylalanine, gamma‐aminobutyric acid, alanine, phenylalanine, tyrosine, glucosamine, guanine, and aspartic acid. Aminoacyl‐tRNA and plant secondary metabolite biosynthesis and amino acid metabolism or synthesis pathways were involved in producing genetic variation under drought conditions. Metabolic changes in light of drought conditions highlighted pools of metabolites that affect the metabolic and physiological adjustment in chickpea that reduced drought impacts.     

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.     

Global change‐type drought‐induced tree mortality: vapor pressure deficit is more important than temperature per se in causing decline in tree health

Drought‐induced tree mortality is occurring across all forested continents and is expected to increase worldwide during the coming century. Regional‐scale forest die‐off influences terrestrial albedo, carbon and water budgets, and land‐surface energy partitioning. Although increased temperatures during drought are widely identified as a critical contributor to exacerbated tree mortality associated with “global‐change‐type drought”, corresponding changes in vapor pressure deficit (D) have rarely been considered explicitly and have not been disaggregated from that of temperature per se. Here, we apply a detailed mechanistic soil–plant–atmosphere model to examine the impacts of drought, increased air temperature (+2°C or +5°C), and increased vapor pressure deficit (D; +1 kPa or +2.5 kPa), singly and in combination, on net primary productivity (NPP) and transpiration and forest responses, especially soil moisture content, leaf water potential, and stomatal conductance. We show that increased D exerts a larger detrimental effect on transpiration and NPP, than increased temperature alone, with or without the imposition of a 3‐month drought. Combined with drought, the effect of increased D on NPP was substantially larger than that of drought plus increased temperature. Thus, the number of days when NPP was zero across the 2‐year simulation was 13 or 14 days in the control and increased temperature scenarios, but increased to approximately 200 days when D was increased. Drought alone increased the number of days of zero NPP to 88, but drought plus increased temperature did not increase the number of days. In contrast, drought and increased D resulted in the number of days when NPP = 0 increasing to 235 (+1 kPa) or 304 days (+2.5 kPa). We conclude that correct identification of the causes of global change‐type mortality events requires explicit consideration of the influence of D as well as its interaction with drought and temperature.     

Role of six European tree species and land‐use legacy for nitrogen and water budgets in forests

Water and nutrient fluxes for single stands of different tree species have been reported in numerous studies, but comparative studies of nutrient and hydrological budgets of common European deciduous tree species are rare. Annual fluxes of water and inorganic nitrogen (N) were established in a 30‐year‐old common garden design with stands of common ash (Fraxinus excelsior), European beech (Fagus sylvatica L.), pedunculate oak (Quercus robur), small‐leaved lime (Tilia cordata Mill.), sycamore maple (Acer pseudoplatanus) and Norway spruce (Picea abies [L.] Karst.) replicated at two sites in Denmark, Mattrup and Vallø during 2 years. Mean annual percolation below the root zone (mm yr^?1±SE, n=4) ranked in the following order: maple (351±38)>lime (284±32), oak (271±25), beech (257±30), ash (307±69)? spruce (75±24). There were few significant tree species effects on N fluxes. However, the annual mean N throughfall flux (kg N ha^?1 yr^?1±SE, n=4) for spruce (28±2) was significantly larger than for maple (12±1), beech (11±1) and oak (9±1) stands but not different from that of lime (15±3). Ash had a low mean annual inorganic N throughfall deposition of 9.1 kg ha^?1, but was only present at Mattrup. Annual mean of inorganic N leaching (kg ha^?1 yr^?1±SE, n=4) did not differ significantly between species despite of contrasting tree species mean values; beech (25±9)>oak (16±10), spruce (15±8), lime (14±8)? maple (1.9±1), ash (2.0±1). The two sites had similar throughfall N fluxes, whereas the annual leaching of N was significantly higher at Mattrup than at Vallø. Accordingly, the sites differed in soil properties in relation to rates and dynamics of N cycling. We conclude that tree species affect the N cycle differently but the legacy of land use exerted a dominant control on the N cycle within the short‐term perspective (30 years) of these stands.     

Elevated [CO2] does not ameliorate the negative effects of elevated temperature on drought‐induced mortality in Eucalyptus radiata seedlings

It has been reported that elevated temperature accelerates the time‐to‐mortality in plants exposed to prolonged drought, while elevated [CO₂] acts as a mitigating factor because it can reduce stomatal conductance and thereby reduce water loss. We examined the interactive effects of elevated [CO₂] and temperature on the inter‐dependent carbon and hydraulic characteristics associated with drought‐induced mortality in Eucalyptus radiata seedlings grown in two [CO₂] (400 and 640 μL L^?1) and two temperature (ambient and ambient +4 °C) treatments. Seedlings were exposed to two controlled drying and rewatering cycles, and then water was withheld until plants died. The extent of xylem cavitation was assessed as loss of stem hydraulic conductivity. Elevated temperature triggered more rapid mortality than ambient temperature through hydraulic failure, and was associated with larger water use, increased drought sensitivities of gas exchange traits and earlier occurrence of xylem cavitation. Elevated [CO₂] had a negligible effect on seedling response to drought, and did not ameliorate the negative effects of elevated temperature on drought. Our findings suggest that elevated temperature and consequent higher vapour pressure deficit, but not elevated [CO₂], may be the primary contributors to drought‐induced seedling mortality under future climates.     

Fire‐induced tree mortality in a neotropical forest: the roles of bark traits,tree size,wood density and fire behavior

Large‐scale wildfires are expected to accelerate forest dieback in Amazônia, but the fire vulnerability of tree species remains uncertain, in part due to the lack of studies relating fire‐induced mortality to both fire behavior and plant traits. To address this gap, we established two sets of experiments in southern Amazonia. First, we tested which bark traits best predict heat transfer rates (R) through bark during experimental bole heating. Second, using data from a large‐scale fire experiment, we tested the effects of tree wood density (WD), size, and estimated R (inverse of cambium insulation) on tree mortality after one to five fires. In the first experiment, bark thickness explained 82% of the variance in R, while the presence of water in the bark reduced the difference in temperature between the heat source and the vascular cambium, perhaps because of high latent heat of vaporization. This novel finding provides an important insight for improving mechanistic models of fire‐induced cambium damage from tropical to temperate regions. In the second experiment, tree mortality increased with increasing fire intensity (i.e. as indicated by bark char height on tree boles), which was higher along the forest edge, during the 2007 drought, and when the fire return interval was 3 years instead of one. Contrary to other tropical studies, the relationship between mortality and fire intensity was strongest in the year following the fires, but continued for 3 years afterwards. Tree mortality was low (≤20%) for thick‐barked individuals (≥18 mm) subjected to medium‐intensity fires, and significantly decreased as a function of increasing tree diameter, height and wood density. Hence, fire‐induced tree mortality was influenced not only by cambium insulation but also by other traits that reduce the indirect effects of fire. These results can be used to improve assessments of fire vulnerability of tropical forests.     

Effects of increasing root carbon investment on the mortality and resprouting of Haloxylon ammodendron seedlings under drought

• Tree mortality induced by drought is one of the most complex processes in ecology. Although two mechanisms associated with water and carbon balance are proposed to explain tree mortality, outstanding problems still exist.
• Here, in order to test how the root system benefits survival and resprouting of Haloxylon ammodendron seedlings, we examined the various water‐ and carbon‐related physiological indicators (shoot water potential, photosynthesis, dark respiration, hydraulic conductance and non‐structural carbohydrates [NSC]) of H. ammodendron seedlings, which were grown in drought and control conditions throughout a grow season in greenhouse.
• The survival time of the seedling root system (died 70 days after drought) doubled the survival time of the shoot (died at 35 days). Difference in survival time between shoot and root resulted from sustained root respiration supported by increased NSC in roots under drought. Furthermore, investment into the root contributed to resprouting following drought. Based on these results, a death criterion is proposed for this species. The time sequence of major events indicated that drought shifted carbon allocation between shoot and root and altered the flux among different sinks (growth, respiration or storage). The interaction of water and carbon processes determined death or survival of droughted H. ammodendron seedlings.
• These findings revealed that the ‘root protection’ strategy is critical in determining survival and resprouting of this species, and provided insights into the effects of carbon and water dynamics on tree mortality.

     

A synthesis of radial growth patterns preceding tree mortality

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

Stem hydraulic capacitance decreases with drought stress: implications for modelling tree hydraulics in the Mediterranean oak Quercus ilex

Hydraulic modelling is a primary tool to predict plant performance in future drier scenarios. However, as most tree models are validated under non‐stress conditions, they may fail when water becomes limiting. To simulate tree hydraulic functioning under moist and dry conditions, the current version of a water flow and storage mechanistic model was further developed by implementing equations that describe variation in xylem hydraulic resistance (R_X) and stem hydraulic capacitance (C_S) with predawn water potential (Ψ_PD). The model was applied in a Mediterranean forest experiencing intense summer drought, where six Quercus ilex trees were instrumented to monitor stem diameter variations and sap flow, concurrently with measurements of predawn and midday leaf water potential. Best model performance was observed when C_S was allowed to decrease with decreasing Ψ_PD. Hydraulic capacitance decreased from 62 to 25 kg m^?3 MPa^?1 across the growing season. In parallel, tree transpiration decreased to a greater extent than the capacitive water release and the contribution of stored water to transpiration increased from 2.0 to 5.1%. Our results demonstrate the importance of stored water and seasonality in C_S for tree hydraulic functioning, and they suggest that C_S should be considered to predict the drought response of trees with models.