Drought causes reduced growth of trembling aspen in western Canada

Adequate and advance knowledge of the response of forest ecosystems to temperature‐induced drought is critical for a comprehensive understanding of the impacts of global climate change on forest ecosystem structure and function. Recent massive decline in aspen‐dominated forests and an increased aspen mortality in boreal forests have been associated with global warming, but it is still uncertain whether the decline and mortality are driven by drought. We used a series of ring‐width chronologies from 40 trembling aspen (Populus tremuloides Michx.) sites along a latitudinal gradient (from 52° to 58°N) in western Canada, in an attempt to clarify the impacts of drought on aspen growth by using Standardized Precipitation Index (SPI) and Standardized Precipitation Evapotranspiration Index (SPEI). Results indicated that prolonged and large‐scale droughts had a strong negative impact on trembling aspen growth. Furthermore, the spatiotemporal variability of drought indices is useful for explaining the spatial heterogeneity in the radial growth of trembling aspen. Due to ongoing global warming and rising temperatures, it is likely that severer droughts with a higher frequency will occur in western Canada. As trembling aspen is sensitive to drought, we suggest that drought indices could be applied to monitor the potential effects of increased drought stress on aspen trees growth, achieve classification of eco‐regions and develop effective mitigation strategies to maintain western Canadian boreal forests.     

Generalist predation on forest tent caterpillar varies with forest stand composition: an experimental study across multiple life stages

1. Generalist enemies can regulate low‐density forest insect populations, and are widely considered to cause greater mortality in more diverse habitats. Forest tent caterpillars (Malacosoma disstria Hübner; FTC) are a major defoliator of aspen (Populus tremuloides Micheaux) in the boreal forest, a region with a mosaic of forest stand types. This heterogeneity may influence FTC outbreaks if generalist predation or parasitism differs among stands of different tree composition. 2. Using exclusion experiments we estimate predation and parasitism of FTC across multiple life‐history stages in low‐density populations occupying both aspen (low diversity) and mixedwood stands (high diversity). 3. Arthropod and avian generalist predators were responsible for most natural enemy‐caused mortality of immature FTC, but their relative impacts varied among FTC life‐history stages. Contrary to expectation, predation on late instar larvae and pupae was higher in the less diverse aspen stands and early instar mortality did not differ. 4. By considering multiple life‐history stages, our results provide a more comprehensive view of natural enemy‐caused morality of immature FTC. Because generalist predation on FTC was higher in aspen than in mixedwood stands, we suggest that FTC populations may be slower to reach outbreak levels in aspen stands.     

Slow lifelong growth predisposes Populus tremuloides trees to mortality

Widespread dieback of aspen forests, sometimes called sudden aspen decline, has been observed throughout much of western North America, with the highest mortality rates in the southwestern United States. Recent aspen mortality has been linked to drought stress and elevated temperatures characteristic of conditions expected under climate change, but the role of individual aspen tree growth patterns in contributing to recent tree mortality is less well known. We used tree-ring data to investigate the relationship between an individual aspen tree’s lifetime growth patterns and mortality. Surviving aspen trees had consistently higher average growth rates for at least 100 years than dead trees. Contrary to observations from late successional species, slow initial growth rates were not associated with a longer lifespan in aspen. Aspen trees that died had slower lifetime growth and slower growth at various stages of their lives than those that survived. Differences in average diameter growth between live and dead trees were significant (α = 0.05) across all time periods tested. Our best logistical model of aspen mortality indicates that younger aspen trees with lower recent growth rates and higher frequencies of abrupt growth declines had an increased risk of mortality. Our findings highlight the need for species-specific mortality functions in forest succession models. Size-dependent mortality functions suitable for late successional species may not be appropriate for species with different life history strategies. For some early successional species, like aspen, slow growth at various stages of the tree’s life is associated with increased mortality risk.     

Drought characteristics' role in widespread aspen forest mortality across Colorado,USA

Globally documented widespread drought‐induced forest mortality has important ramifications for plant community structure, ecosystem function, and the ecosystem services provided by forests. Yet the characteristics of drought seasonality, severity, and duration that trigger mortality events have received little attention despite evidence of changing precipitation regimes, shifting snow melt timing, and increasing temperature stress. This study draws upon stand level ecohydrology and statewide climate and spatial analysis to examine the drought characteristics implicated in the recent widespread mortality of trembling aspen (Populus tremuloides Michx.). We used isotopic observations of aspen xylem sap to determine water source use during natural and experimental drought in a region that experienced high tree mortality. We then drew upon multiple sources of climate data to characterize the drought that triggered aspen mortality. Finally, regression analysis was used to examine the drought characteristics most associated with the spatial patterns of aspen mortality across Colorado. Isotopic analysis indicated that aspens generally utilize shallow soil moisture with little plasticity during drought stress. Climate analysis showed that the mortality‐inciting drought was unprecedented in the observational record, especially in 2002 growing season temperature and evaporative deficit, resulting in record low shallow soil moisture reserves. High 2002 summer temperature and low shallow soil moisture were most associated with the spatial patterns of aspen mortality. These results suggest that the 2002 drought subjected Colorado aspens to the most extreme growing season water stress of the past century by creating high atmospheric moisture demand and depleting the shallow soil moisture upon which aspens rely. Our findings highlight the important role of drought characteristics in mediating widespread aspen forest mortality, link this aspen die‐off to regional climate change trends, and provide insight into future climate vulnerability of these forests.     

Massive mortality of aspen following severe drought along the southern edge of the Canadian boreal forest

Drought‐induced, regional‐scale dieback of forests has emerged as a global concern that is expected to escalate under model projections of climate change. Since 2000, drought of unusual severity, extent, and duration has affected large areas of western North America, leading to regional‐scale dieback of forests in the southwestern US. We report on drought impacts on forests in a region farther north, encompassing the transition between boreal forest and prairie in western Canada. A central question is the significance of drought as an agent of large‐scale tree mortality and its potential future impact on carbon cycling in this cold region. We used a combination of plot‐based, meteorological, and remote sensing measures to map and quantify aboveground, dead biomass of trembling aspen (Populus tremuloides Michx.) across an 11.5 Mha survey area where drought was exceptionally severe during 2001–2002. Within this area, a satellite‐based land cover map showed that aspen‐dominated broadleaf forests occupied 2.3 Mha. Aerial surveys revealed extensive patches of severe mortality (>55%) resembling the impacts of fire. Dead aboveground biomass was estimated at 45 Mt, representing 20% of the total aboveground biomass, based on a spatial interpolation of plot‐based measurements. Spatial variation in percentage dead biomass showed a moderately strong correlation with drought severity. In the prairie‐like, southern half of the study area where the drought was most severe, 35% of aspen biomass was dead, compared with an estimated 7% dead biomass in the absence of drought. Drought led to an estimated 29 Mt increase in dead biomass across the survey area, corresponding to 14 Mt of potential future carbon emissions following decomposition. Many recent, comparable episodes of drought‐induced forest dieback have been reported from around the world, which points to an emerging need for multiscale monitoring approaches to quantify drought effects on woody biomass and carbon cycling across large areas.     

Drought's legacy: multiyear hydraulic deterioration underlies widespread aspen forest die‐off and portends increased future risk

Forest mortality constitutes a major uncertainty in projections of climate impacts on terrestrial ecosystems and carbon‐cycle feedbacks. Recent drought‐induced, widespread forest die‐offs highlight that climate change could accelerate forest mortality with its diverse and potentially severe consequences for the global carbon cycle, ecosystem services, and biodiversity. How trees die during drought over multiple years remains largely unknown and precludes mechanistic modeling and prediction of forest die‐off with climate change. Here, we examine the physiological basis of a recent multiyear widespread die‐off of trembling aspen (Populus tremuloides) across much of western North America. Using observations from both native trees while they are dying and a rainfall exclusion experiment on mature trees, we measure hydraulic performance over multiple seasons and years and assess pathways of accumulated hydraulic damage. We test whether accumulated hydraulic damage can predict the probability of tree survival over 2 years. We find that hydraulic damage persisted and increased in dying trees over multiple years and exhibited few signs of repair. This accumulated hydraulic deterioration is largely mediated by increased vulnerability to cavitation, a process known as cavitation fatigue. Furthermore, this hydraulic damage predicts the probability of interyear stem mortality. Contrary to the expectation that surviving trees have weathered severe drought, the hydraulic deterioration demonstrated here reveals that surviving regions of these forests are actually more vulnerable to future droughts due to accumulated xylem damage. As the most widespread tree species in North America, increasing vulnerability to drought in these forests has important ramifications for ecosystem stability, biodiversity, and ecosystem carbon balance. Our results provide a foundation for incorporating accumulated drought impacts into climate–vegetation models. Finally, our findings highlight the critical role of drought stress accumulation and repair of stress‐induced damage for avoiding plant mortality, presenting a dynamic and contingent framework for drought impacts on forest ecosystems.     

Experimental climate warming alters aspen and birch phytochemistry and performance traits for an outbreak insect herbivore

Climate change and insect outbreaks are key factors contributing to regional and global patterns of increased tree mortality. While links between these environmental stressors have been established, our understanding of the mechanisms by which elevated temperature may affect tree–insect interactions is limited. Using a forest warming mesocosm, we investigated the influence of elevated temperature on phytochemistry, tree resistance traits, and insect performance. Specifically, we examined warming effects on forest tent caterpillar (Malacosoma disstria) and host trees aspen (Populus tremuloides) and birch (Betula papyrifera). Trees were grown under one of three temperature treatments (ambient, +1.7 °C, +3.4 °C) in a multiyear open‐air warming experiment. In the third and fourth years of warming (2011, 2012), we assessed foliar nutrients and defense chemistry. Elevated temperatures altered foliar nitrogen, carbohydrates, lignin, and condensed tannins, with differences in responses between species and years. In 2012, we performed bioassays using a common environment approach to evaluate plant‐mediated indirect warming effects on larval performance. Warming resulted in decreased food conversion efficiency and increased consumption, ultimately with minimal effect on larval development and biomass. These changes suggest that insects exhibited compensatory feeding due to reduced host quality. Within the context of observed phytochemical variation, primary metabolites were stronger predictors of insect performance than secondary metabolites. Between‐year differences in phytochemical shifts corresponded with substantially different weather conditions during these two years. By sampling across years within an ecologically realistic and environmentally open setting, our study demonstrates that plant and insect responses to warming can be temporally variable and context dependent. Results indicate that elevated temperatures can alter phytochemistry, tree resistance traits, and herbivore feeding, but that annual weather variability may modulate warming effects leading to uncertain consequences for plant–insect interactions with projected climate change.     

Mortality and growth of trees in peat-swamp and heath forests in Central Kalimantan after severe drought

Lowland forests in Central Kalimantan, Indonesian Borneo, are endangered by land conversion and the increasing frequency of severe drought. Knowledge of the tolerance of tropical trees to drought is urgent for the management of these lowland habitats. The short-term effects of drought on tree demography (mortality and growth) were investigated in an ever-wet riparian peat-swamp forest and a heath forest on coarse sandy soil after the 1997 El Niño Southern Oscillation (ENSO) event. This drought was unusually severe because little rain fell during the following rainy season. However, forest-wide mortality following the drought (1997–1999) was not critically high in the peat-swamp (6.13% yr^?1) or heath (4.26% yr^?1) forest. In both forests, standing trees frequently died during the dry season following the drought. The riparian peat-swamp forest was not flooded until 1998, after the prolonged drought in 1997. The hummock–hollow microtopography resulted in differential mortality of peat-swamp trees. On tall hummocks, standing death increased two-fold (4.99% yr^?1) during the dry season, whereas uprooting decreased by one-third (0.85% yr^?1) during the following rainy season. In contrast, tree growth was not affected by hummock height. Common canopy species were concentrated on tall hummocks and died standing more often than did understory species found in hollows, indicating species-specific mortality after the drought. The large stand basal area relative to the forest-wide growth rate in diameter suggested less resilience to drought by peat-swamp (45.6 m² ha^?1 and 0.0186 ln[cm] yr^?1) than heath (27.9 m² ha^?1 and 0.0232 ln[cm] yr^?1) forest. A single severe drought did not cause dramatic changes in the peat-swamp and heath forests; however, an increasing frequency of droughts similar in severity to that of the 1997 ENSO event may have the potential to alter the community structure and dynamics, leading to a consistent decline in Bornean lowland forests.     

Differential declines in Alaskan boreal forest vitality related to climate and competition

Rapid warming and changes in water availability at high latitudes alter resource abundance, tree competition, and disturbance regimes. While these changes are expected to disrupt the functioning of boreal forests, their ultimate implications for forest composition are uncertain. In particular, recent site‐level studies of the Alaskan boreal forest have reported both increases and decreases in productivity over the past few decades. Here, we test the idea that variations in Alaskan forest growth and mortality rates are contingent on species composition. Using forest inventory measurements and climate data from plots located throughout interior and south‐central Alaska, we show significant growth and mortality responses associated with competition, midsummer vapor pressure deficit, and increased growing season length. The governing climate and competition processes differed substantially across species. Surprisingly, the most dramatic climate response occurred in the drought tolerant angiosperm species, trembling aspen, and linked high midsummer vapor pressure deficits to decreased growth and increased insect‐related mortality. Given that species composition in the Alaskan and western Canadian boreal forests is projected to shift toward early‐successional angiosperm species due to fire regime, these results underscore the potential for a reduction in boreal productivity stemming from increases in midsummer evaporative demand.     

Large drought‐induced aboveground live biomass losses in southern Rocky Mountain aspen forests

Widespread drought‐induced forest mortality has been documented across multiple tree species in North America in recent decades, but it is a poorly understood component in terrestrial carbon (C) budgets. Recent severe drought in concert with elevated temperature likely triggered widespread forest mortality of trembling aspen (Populus tremuloides), the most widely distributed tree species in North America. The impact on the regional C budgets and spatial pattern of this drought‐induced tree mortality, which has been termed ‘sudden aspen decline (SAD)’, is not well known and could contribute to increased regional C emissions, an amplifying feedback to climate change. We conducted a regional assessment of drought‐induced live aboveground biomass (AGB) loss from SAD across 915 km² of southwestern Colorado, USA, and investigated the influence of topography on the severity of mortality by combining field measures, remotely sensed nonphotosynthetically active vegetation and a digital elevation model. Mean [± standard deviation (SD)] remote sensing estimate of live AGB loss was 60.3 ± 37.3 Mg ha^?1, which was 30.7% of field measured AGB, totaling 2.7 Tg of potential C emissions from this dieback event. Aspen forest health could be generally categorized as healthy (0–30% field measured canopy dieback), intermediate (31–50%), and SAD (51–100%), with the remote sensing estimated mean (± SD) live AGB losses of 26.4 ± 15.1, 64.5 ± 9.2, and 108.5 ± 24.0 Mg ha^?1, respectively. There was a pronounced clustering pattern of SAD on south‐facing slopes due to relatively drier and warmer conditions, but no apparent spatial gradient was found for elevation and slope. This study demonstrates the feasibility of utilizing remote sensing to assess the ramification of climate‐induced forest mortality on ecosystems and suggests promising opportunities for systematic large‐scale C dynamics monitoring of tree dieback, which would improve estimates of C budgets of North America with climate change.     

Factors affecting fall down rates of dead aspen (Populus tremuloides) biomass following severe drought in west‐central Canada

Increases in mortality of trembling aspen (Populus tremuloides Michx.) have been recorded across large areas of western North America following recent periods of exceptionally severe drought. The resultant increase in standing, dead tree biomass represents a significant potential source of carbon emissions to the atmosphere, but the timing of emissions is partially driven by dead‐wood dynamics which include the fall down and breakage of dead aspen stems. The rate at which dead trees fall to the ground also strongly influences the period over which forest dieback episodes can be detected by aerial surveys or satellite remote sensing observations. Over a 12‐year period (2000–2012), we monitored the annual status of 1010 aspen trees that died during and following a severe regional drought within 25 study areas across west‐central Canada. Observations of stem fall down and breakage (snapping) were used to estimate woody biomass transfer from standing to downed dead wood as a function of years since tree death. For the region as a whole, we estimated that >80% of standing dead aspen biomass had fallen after 10 years. Overall, the rate of fall down was minimal during the year following stem death, but thereafter fall rates followed a negative exponential equation with k = 0.20 per year. However, there was high between‐site variation in the rate of fall down (k = 0.08–0.37 per year). The analysis showed that fall down rates were positively correlated with stand age, site windiness, and the incidence of decay fungi (Phellinus tremulae (Bond.) Bond. and Boris.) and wood‐boring insects. These factors are thus likely to influence the rate of carbon emissions from dead trees following periods of climate‐related forest die‐off episodes.     

Evaluation of climate‐related carbon turnover processes in global vegetation models for boreal and temperate forests

Turnover concepts in state‐of‐the‐art global vegetation models (GVMs) account for various processes, but are often highly simplified and may not include an adequate representation of the dominant processes that shape vegetation carbon turnover rates in real forest ecosystems at a large spatial scale. Here, we evaluate vegetation carbon turnover processes in GVMs participating in the Inter‐Sectoral Impact Model Intercomparison Project (ISI‐MIP, including HYBRID4, JeDi, JULES, LPJml, ORCHIDEE, SDGVM, and VISIT) using estimates of vegetation carbon turnover rate (k) derived from a combination of remote sensing based products of biomass and net primary production (NPP). We find that current model limitations lead to considerable biases in the simulated biomass and in k (severe underestimations by all models except JeDi and VISIT compared to observation‐based average k), likely contributing to underestimation of positive feedbacks of the northern forest carbon balance to climate change caused by changes in forest mortality. A need for improved turnover concepts related to frost damage, drought, and insect outbreaks to better reproduce observation‐based spatial patterns in k is identified. As direct frost damage effects on mortality are usually not accounted for in these GVMs, simulated relationships between k and winter length in boreal forests are not consistent between different regions and strongly biased compared to the observation‐based relationships. Some models show a response of k to drought in temperate forests as a result of impacts of water availability on NPP, growth efficiency or carbon balance dependent mortality as well as soil or litter moisture effects on leaf turnover or fire. However, further direct drought effects such as carbon starvation (only in HYBRID4) or hydraulic failure are usually not taken into account by the investigated GVMs. While they are considered dominant large‐scale mortality agents, mortality mechanisms related to insects and pathogens are not explicitly treated in these models.     

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.     

Growth and mortality of trembling aspen (Populus tremuloides) in response to artificial defoliation

To simulate the effects of forest tent caterpillar (FTC) defoliation on trembling aspen growth and mortality, an artificial defoliation experiment was performed over three years in young aspen stands of northwestern Quebec. Defoliation plots of 15 × 15 m were established on three sites, together with associated control stands of pure trembling aspen. In 2007, root collar diameters were measured and positions of all trees were mapped prior defoliation. Severe FTC defoliation was simulated for three successive years (2007–2009) by manually removing all leaves from all but 7–10% of the trees present in the defoliation plots. Yearly surveys of growth and mortality were conducted until 2010 to evaluate defoliation effects on defoliated as well as surrounding undefoliated trees. In absence of other factors, growth and mortality of trembling aspen decreased and increased, respectively, after defoliation. Our study further revealed that small diameter trees died after one year of artificial defoliation, while larger-diameter trees died after repeated defoliations. Distributions of tree mortality tended to be aggregated at small scales (<5 m), corroborating gap patterns observed in mature stands following FTC outbreaks. This experiment revealed that trembling aspen mortality can be directly attributed solely to defoliation. Repeated defoliations during FTC outbreaks have the potential to profoundly modify stand productivity and structure by reducing tree growth and increasing tree mortality in the absence of predisposing factors.     

Drought induces lagged tree mortality in a subalpine forest in the Rocky Mountains

Extreme climatic events are key factors in initiating gradual or sudden changes in forest ecosystems through the promotion of severe, tree-killing disturbances such as fire, blowdown, and widespread insect outbreaks. In contrast to these climatically-incited disturbances, little is known about the more direct effect of drought on tree mortality, especially in high-elevation forests. Therefore projections of drought-induced mortality under future climatic conditions remain uncertain. For a subalpine forest landscape in the Rocky Mountains of northern Colorado (USA), we quantified lag effects of drought on mortality of Engelmann spruce Picea engelmannii , subalpine fir Abies lasiocarpa , and lodgepole pine Pinus contorta . For the period 1910–2004, we related death dates of 164 crossdated dead trees to early-season and late-season droughts. Following early-season droughts, spruce mortality increased over five years and fir mortality increased sharply over 11 years. Following late-season droughts, spruce showed a small increase in mortality within one year, whereas fir showed a consistent period of increased mortality over two years. Pine mortality was not affected by drought. Low pre-drought radial growth rates predisposed spruce and fir to drought-related mortality. Spruce and fir trees that died during a recent drought (2000–2004) had significantly lower pre-drought growth rates than live neighbour trees. Overall, we found large interspecific differences in drought-related mortality with fir showing the strongest effect followed by spruce and pine. This direct influence of climatic variability on differential tree mortality has the potential for driving large-scale changes in subalpine forests of the Rocky Mountains.     

Insect‐induced tree mortality of boreal forests in eastern Canada under a changing climate

Forest insects are major disturbances that induce tree mortality in eastern coniferous (or fir-spruce) forests in eastern North America. The spruce budworm (SBW) (Choristoneura fumiferana [Clemens]) is the most devastating insect causing tree mortality. However, the relative importance of insect-caused mortality versus tree mortality caused by other agents and how this relationship will change with climate change is not known. Based on permanent sample plots across eastern Canada, we combined a logistic model with a negative model to estimate tree mortality. The results showed that tree mortality increased mainly due to forest insects. The mean difference in annual tree mortality between plots disturbed by insects and those without insect disturbance was 0.0680 per year (P < 0.0001, T-test), and the carbon sink loss was about 2.87t C ha⁻¹ year⁻¹ larger than in natural forests. We also found that annual tree mortality increased significantly with the annual climate moisture index (CMI) and decreased significantly with annual minimum temperature (T_min), annual mean temperature (T_mean) and the number of degree days below 0°C (DD0), which was inconsistent with previous studies (Adams et al. 2009; van Mantgem et al. 2009; Allen et al. 2010). Furthermore, the results for the trends in the magnitude of forest insect outbreaks were consistent with those of climate factors for annual tree mortality. Our results demonstrate that forest insects are the dominant cause of the tree mortality in eastern Canada but that tree mortality induced by insect outbreaks will decrease in eastern Canada under warming climate.     

Long-term drought effects on the thermal sensitivity of Amazon forest trees

The continued functioning of tropical forests under climate change depends on their resilience to drought and heat. However, there is little understanding of how tropical forests will respond to combinations of these stresses, and no field studies to date have explicitly evaluated whether sustained drought alters sensitivity to temperature. We measured the temperature response of net photosynthesis, foliar respiration and the maximum quantum efficiency of photosystem II (F_v/F_m) of eight hyper-dominant Amazonian tree species at the world's longest-running tropical forest drought experiment, to investigate the effect of drought on forest thermal sensitivity. Despite a 0.6°C–2°C increase in canopy air temperatures following long-term drought, no change in overall thermal sensitivity of net photosynthesis or respiration was observed. However, photosystem II tolerance to extreme-heat damage (T₅₀) was reduced from 50.0 ± 0.3°C to 48.5 ± 0.3°C under drought. Our results suggest that long-term reductions in precipitation, as projected across much of Amazonia by climate models, are unlikely to greatly alter the response of tropical forests to rising mean temperatures but may increase the risk of leaf thermal damage during heatwaves.     

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,...     

Forest Fragmentation Alters the Population Dynamics of a Late‐successional Tropical Tree

Tropical late‐successional tree species are at high risk of local extinction due to habitat loss and fragmentation. Population‐growth rates in fragmented populations are predicted to decline as a result of reduced fecundity, survival and growth. We examined the demographic effects of habitat fragmentation by comparing the population dynamics of the late‐successional tree Poulsenia armata (Moraceae) in southern Mexico between a continuous forest and several forest fragments using integral projection models (IPMs) during 2010–2012. Forest fragmentation did not lead to differences in population density and even resulted in a higher population‐growth rate (λ) in fragments compared to continuous forests. Habitat fragmentation had drastic effects on the dynamics of P. armata, causing the population structure to shift toward smaller sizes. Fragmented populations experienced a significant decrease in juvenile survival and growth compared to unaltered populations. Adult survival and growth made the greatest relative contributions to λ in both habitat types during 2011–2012. However, the relative importance of juvenile survival and growth to λ was highest in the fragmented forest in 2010–2011. Our Life Table Response Experiment analysis revealed that positive contributions of adult fecundity explained most of the variation of λ between both habitats and annual periods. Finally, P. armata has a relatively slow speed of recovery after disturbances, compromising persistence of fragmented populations. Developing a mechanistic understanding of how forest fragmentation affects plant population dynamics, as done here, will prove essential for the preservation of natural areas.     

Stability in the patterns of long‐term development and growth of the Canadian spruce–moss forest

Aim The spruce–moss forest is the main forest ecosystem of the North American boreal forest. We used stand structure and fire data to examine the long‐term development and growth of the spruce–moss ecosystem. We evaluate the stability of the forest with time and the conditions needed for the continuing regeneration, growth and re‐establishment of black spruce (Picea mariana) trees. Location The study area occurs in Québec, Canada, and extends from 70°00′ to 72°00′ W and 47°30′ to 56°00′ N. Methods A spatial inventory of spruce–moss forest stands was performed along 34 transects. Nineteen spruce–moss forests were selected. A 500 m² quadrat at each site was used for radiocarbon and tree‐ring dating of time since last fire (TSLF). Size structure and tree regeneration in each stand were described based on diameter distribution of the dominant and co‐dominant tree species [black spruce and balsam fir (Abies balsamea)]. Results The TSLF of the studied forests ranges from 118 to 4870 cal. yr bp . Forests < 325 cal. yr bp are dominated by trees of the first post‐fire cohort and are not yet at equilibrium, whereas older forests show a reverse‐J diameter distribution typical of mature, old‐growth stands. The younger forests display faster height and radial growth‐rate patterns than the older forests, due to factors associated with long‐term forest development. Each of the stands examined established after severe fires that consumed all the soil organic material. Main conclusions Spruce–moss forests are able to self‐regenerate after fires that consume the organic layer, thus allowing seed regeneration at the soil surface. In the absence of fire the forests can remain in an equilibrium state. Once the forests mature, tree productivity eventually levels off and becomes stable. Further proof of the enduring stability of these forests, in between fire periods, lies in the ages of the stands. Stands with a TSLF of 325–4870 cal. yr bp all exhibited the same stand structure, tree growth rates and species characteristics. In the absence of fire, the spruce–moss forests are able to maintain themselves for thousands of years with no apparent degradation or change in forest type.