Variation in woody plant mortality and dieback from severe drought among soils, plant groups, and species within a northern Arizona ecotone

Vegetation change from drought-induced mortality can alter ecosystem community structure, biodiversity, and services. Although drought-induced mortality of woody plants has increased globally with recent warming, influences of soil type, tree and shrub groups, and species are poorly understood. Following the severe 2002 drought in northern Arizona, we surveyed woody plant mortality and canopy dieback of live trees and shrubs at the forest–woodland ecotone on soils derived from three soil parent materials (cinder, flow basalt, sedimentary) that differed in texture and rockiness. Our first of three major findings was that soil parent material had little effect on mortality of both trees and shrubs, yet canopy dieback of trees was influenced by parent material; dieback was highest on the cinder for pinyon pine (Pinus edulis) and one-seed juniper (Juniperus monosperma). Ponderosa pine (Pinus ponderosa) dieback was not sensitive to parent material. Second, shrubs had similar mortality, but greater canopy dieback, than trees. Third, pinyon and ponderosa pines had greater mortality than juniper, yet juniper had greater dieback, reflecting different hydraulic characteristics among these tree species. Our results show that impacts of severe drought on woody plants differed among tree species and tree and shrub groups, and such impacts were widespread over different soils in the southwestern U.S. Increasing frequency of severe drought with climate warming will likely cause similar mortality to trees and shrubs over major soil types at the forest–woodland ecotone in this region, but due to greater mortality of other tree species, tree cover will shift from a mixture of species to dominance by junipers and shrubs. Surviving junipers and shrubs will also likely have diminished leaf area due to canopy dieback.     

Measuring canopy loss and climatic thresholds from an extreme drought along a fivefold precipitation gradient across Texas

Globally, trees are increasingly dying from extreme drought, a trend that is expected to increase with climate change. Loss of trees has significant ecological, biophysical, and biogeochemical consequences. In 2011, a record drought caused widespread tree mortality in Texas. Using remotely sensed imagery, we quantified canopy loss during and after the drought across the state at 30‐m spatial resolution, from the eastern pine/hardwood forests to the western shrublands, a region that includes the boundaries of many species ranges. Canopy loss observations in ~200 multitemporal fine‐scale orthophotos (1‐m) were used to train coarser Landsat imagery (30‐m) to create 30‐m binary statewide canopy loss maps. We found that canopy loss occurred across all major ecoregions of Texas, with an average loss of 9.5%. The drought had the highest impact in post oak woodlands, pinyon‐juniper shrublands and Ashe juniper woodlands. Focusing on a 100‐km by ~1,000‐km transect spanning the State's fivefold east–west precipitation gradient (~1,500 to ~300 mm), we compared spatially explicit 2011 climatic anomalies to our canopy loss maps. Much of the canopy loss occurred in areas that passed specific climatic thresholds: warm season anomalies in mean temperature (+1.6°C) and vapor pressure deficit (VPD, +0.66 kPa), annual percent deviation in precipitation (?38%), and 2011 difference between precipitation and potential evapotranspiration (?1,206 mm). Although similarly low precipitation occurred during the landmark 1950s drought, the VPD and temperature anomalies observed in 2011 were even greater. Furthermore, future climate data under the representative concentration pathway 8.5 trajectory project that average values will surpass the 2011 VPD anomaly during the 2070–2099 period and the temperature anomaly during the 2040–2099 period. Identifying vulnerable ecological systems to drought stress and climate thresholds associated with canopy loss will aid in predicting how forests will respond to a changing climate and how ecological landscapes will change in the near term.     

Multi-scale species density model for conserving an endangered songbird

Habitat modeling across a landscape that has gradients of habitat conditions requires potential predictor data that can be quantified at biologically relevant scales. We used remotely sensed data to develop a multi-scale density model in 2018 for the golden-cheeked warbler (Setophaga chrysoparia; warbler), a species that breeds in Ashe juniper (Juniperus ashei)-oak (Quercus spp.) woodlands in central Texas, USA. We first classified Ashe juniper and broadleaf tree cover at a 1-m resolution and used this to map potential habitat across the warbler's >67,000-km² breeding range. We then designed a survey for estimating warbler density based on hierarchical distance sampling. We used stratified random sampling to survey for male warblers at 1,804 points across the continuum of tree canopy cover and composition and detected 810 warblers during our surveys. We developed a suite of potential predictor variables for modeling warbler density that reflected vegetation, topography, climate, and anthropogenic land use conditions across the breeding range and developed these at 3 scales representing the territory, site, and landscape. We modeled warbler density and used the best fit model to produce a spatially explicit estimate. Predicted warbler density was influenced by tree canopy cover and canopy height at the territory scale (100-m radius); tree canopy cover, percent of the canopy comprised of juniper, and an interaction between canopy cover and compound topographic index at the site scale (1-km radius); and annual temperature range at the landscape scale (5-km radius). We estimated a population size of 217,444 male warblers (95% CI = 153,917–311,965) and >3,000 males in each recovery unit. After controlling for the duration of point count surveys, our estimate of population size was similar to that reported from the only previous breeding range survey conducted in 2008–2009. Our model results indicated that management activities to increase warbler density should promote woodlands with high tree canopy cover, approximately 60–80% Ashe juniper composition, and tree heights >3 m. In contrast to a patch-based approach, our treatment of habitat variables as continuous helped to credibly map the warbler distribution across areas with broad transitions from woodlands to shrublands. By measuring these predictor variables at biologically relevant scales, we allowed the warbler survey data to define habitat relationships instead of using anthropogenically defined habitat patches. Outcomes from our study show the benefits of developing spatial products tailored to individual species of interest for conservation and management decisions.     

Influence of Juniper on Montezuma Quail Habitat Use in Texas

Montezuma quail (Cyrtonyx montezumae) inhabit oak (Quercus spp.)-juniper (Juniperus spp.) woodlands throughout Mexico and the southwestern United States. In Texas, USA, Montezuma quail occur in the Edwards Plateau and Trans-Pecos Mountains and Basins (Trans-Pecos), 2 ecoregions with contrasting juniper patterns. Ashe juniper (Juniperus ashei) dominates in the Edwards Plateau and has been increasing over decades, whereas alligator juniper (Juniperus deppeana) is a co-dominant in the Trans-Pecos and appears to have stable Montezuma quail populations. Our objectives were to compare between ecoregions the relative abundance and habitat use of Montezuma quail in relation to juniper and quantify the influence of juniper on key features of Montezuma quail habitat (grass height, grass cover, forb cover, and forb species richness). We conducted a study from March–August 2018‒2020 in the Edwards Plateau (Kinney and Edwards counties) and Trans-Pecos ecoregions (Jeff Davis County) to evaluate these objectives. We conducted call-back surveys to estimate relative abundance (number of detections/hr) of Montezuma quail and identify used locations. We collected vegetation data at a micro-scale (16-m) at used (n = 32–30 points) and random locations (n = 70–60 points) in each ecoregion. Relative abundance of Montezuma quail was considerably lower in the Edwards Plateau (0.06 ± 0.01 detections/survey hr; ± SE) than in the Trans-Pecos (1.10 ± 0.30 detections/survey hr). In addition, Montezuma quail selected areas of low Ashe juniper cover (<23% cover), density (<7 trees/80 m²), and height (<2 m) in the Edwards Plateau but selected areas of high alligator juniper cover (>18% cover), density (>4 trees/80 m²), and height (2–8 m) in the Trans-Pecos. Moreover, Ashe juniper cover had a significant, negative influence on herbaceous features, whereas alligator juniper exerted little to no influence. Our results suggest that these 2 juniper species have contrasting effects on Montezuma quail space use and their habitat, possibly as a result of the contrasting influence of these junipers on herbaceous understory. Creating habitat for Montezuma quail in the Edwards Plateau generally will involve the reduction of Ashe juniper and creating patches that possess juniper trees of small stature (<2 m) with low amounts of cover (<23%) and density (<7 trees/80 m²), whereas management in the Trans-Pecos will require site-specific assessments based on current alligator juniper influence. © 2021 The Wildlife Society.     

Hydraulic responses to extreme drought conditions in three co-dominant tree species in shallow soil over bedrock

An important component of the hydrological niche involves the partitioning of water sources, but in landscapes characterized by shallow soils over fractured bedrock, root growth is highly constrained. We conducted a study to determine how physical constraints in the root zone affected the water use of three tree species that commonly coexist on the Edwards Plateau of central Texas; cedar elm (Ulmus crassifolia), live oak (Quercus fusiformis), and Ashe juniper (Juniperus ashei). The year of the study was unusually dry; minimum predawn water potentials measured in August were ?8 MPa in juniper, less than ?8 MPa in elm, and ?5 MPa in oak. All year long, species used nearly identical water sources, based on stable isotope analysis of stem water. Sap flow velocities began to decline simultaneously in May, but the rate of decline was fastest for oak and slowest for juniper. Thus, species partitioned water by time when they could not partition water by source. Juniper lost 15–30 % of its stem hydraulic conductivity, while percent loss for oak was 70–75 %, and 90 % for elm. There was no tree mortality in the year of the study, but 2 years later, after an even more severe drought in 2011, we recorded 34, 14, 6, and 1 % mortality among oak, elm, juniper, and Texas persimmon (Diospyros texana), respectively. Among the study species, mortality rates ranked in the same order as the rate of sap flow decline in 2009. Among the angiosperms, mortality rates correlated with wood density, lending further support to the hypothesis that species with more cavitation-resistant xylem are more susceptible to catastrophic hydraulic failure under acute drought.     

Ecological thresholds at the savanna-forest boundary: how plant traits, resources and fire govern the distribution of tropical biomes

Fire shapes the distribution of savanna and forest through complex interactions involving climate, resources and species traits. Based on data from central Brazil, we propose that these interactions are governed by two critical thresholds. The fire-resistance threshold is reached when individual trees have accumulated sufficient bark to avoid stem death, whereas the fire-suppression threshold is reached when an ecosystem has sufficient canopy cover to suppress fire by excluding grasses. Surpassing either threshold is dependent upon long fire-free intervals, which are rare in mesic savanna. On high-resource sites, the thresholds are reached quickly, increasing the probability that savanna switches to forest, whereas low-resource sites are likely to remain as savanna even if fire is infrequent. Species traits influence both thresholds; saplings of savanna trees accumulate bark thickness more quickly than forest trees, and are more likely to become fire resistant during fire-free intervals. Forest trees accumulate leaf area more rapidly than savanna trees, thereby accelerating the transition to forest. Thus, multiple factors interact with fire to determine the distribution of savanna and forest by influencing the time needed to reach these thresholds. Future work should decipher multiple environmental controls over the rates of tree growth and canopy closure in savanna.     

Tree mortality across biomes is promoted by drought intensity,lower wood density and higher specific leaf area

Drought events are increasing globally, and reports of consequent forest mortality are widespread. However, due to a lack of a quantitative global synthesis, it is still not clear whether drought‐induced mortality rates differ among global biomes and whether functional traits influence the risk of drought‐induced mortality. To address these uncertainties, we performed a global meta‐analysis of 58 studies of drought‐induced forest mortality. Mortality rates were modelled as a function of drought, temperature, biomes, phylogenetic and functional groups and functional traits. We identified a consistent global‐scale response, where mortality increased with drought severity [log mortality (trees trees^?1 year^?1) increased 0.46 (95% CI = 0.2–0.7) with one SPEI unit drought intensity]. We found no significant differences in the magnitude of the response depending on forest biomes or between angiosperms and gymnosperms or evergreen and deciduous tree species. Functional traits explained some of the variation in drought responses between species (i.e. increased from 30 to 37% when wood density and specific leaf area were included). Tree species with denser wood and lower specific leaf area showed lower mortality responses. Our results illustrate the value of functional traits for understanding patterns of drought‐induced tree mortality and suggest that mortality could become increasingly widespread in the future.     

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.     

Long‐term mortality patterns of the deep‐rooted Acacia erioloba: The middle class shall die!

Question: Is there a relationship between size and death in the long‐lived, deep‐rooted tree, Acacia erioloba, in a semi‐arid savanna? What is the size‐class distribution of A. erioloba mortality? Does the mortality distribution differ from total tree size distribution? Does A. erioloba mortality distribution match the mortality distributions recorded thus far in other environments? Location: Dronfield Ranch, near Kimberley, Kalahari, South Africa. Methods: A combination of aerial photographs and a satellite image covering 61 year was used to provide long‐term spatial data on mortality. We used aerial photographs of the study area from 1940, 1964, 1984, 1993 and a satellite image from 2001 to follow three plots covering 510 ha. We were able to identify and individually follow ca. 3000 individual trees from 1940 till 2001. Results: The total number of trees increased over time. No relationship between total number of trees and mean tree size was detected. There were no trends over time in total number of deaths per plot or in size distributions of dead trees. Kolmogorov‐Smirnov tests showed no differences in size class distributions for living trees through time. The size distribution of dead trees was significantly different from the size distribution of all trees present on the plots. Overall, the number of dead trees was low in small size classes, reached a peak value when canopy area was 20 ‐ 30 m², and declined in larger size‐classes. Mortality as a ratio of dead vs. total trees peaked at intermediate canopy sizes too. Conclusion: A. erioloba mortality was size‐dependent, peaking at intermediate sizes. The mortality distribution differs from all other tree mortality distributions recorded thus far. We suggest that a possible mechanism for this unusual mortality distribution is intraspecific competition for water in this semi‐arid environment.     

Variable rainfall has a greater effect than fire on the demography of the dominant tree in a semi‐arid Eucalyptus savanna

Rainfall, fire and competition are emphasized as determinants of the density and basal area of woody vegetation in savanna. The semi‐arid savannas of Australia have substantial multi‐year rainfall deficits and insufficient grass fuel to carry annual fire in contrast to the mesic savannas in more northern regions. This study investigates the influence of rainfall deficit and excess, fire and woody competition on the population dynamics of a dominant tree in a semi‐arid savanna. All individuals of Eucalyptus melanophloia were mapped and monitored in three, 1‐ha plots over an 8.5 year period encompassing wet and dry periods. The plots were unburnt, burnt once and burnt twice. A competition index incorporating the size and distance of neighbours to target individuals was determined. Supplementary studies examined seedling recruitment and the transition of juvenile trees into the sapling layer. Mortality of burnt seedlings was related to lignotuber area but the majority of seedlings are fire resistant within 12 months of germination. Most of the juveniles (≤1 cm dbh) of E. melanophloia either died in the dry period or persisted as juveniles throughout 8.5 years of monitoring. Mortality of juveniles was positively related to woody competition and was higher in the dry period than the wet period. The transition of juveniles to a larger size class occurred at extremely low rates, and a subsidiary study along a clearing boundary suggests release from woody competition allows transition into the sapling layer. From three fires the highest proportion of saplings (1–10 cm dbh) reduced to juveniles was only 5.6% suggesting rates of ‘top‐kill’ of E. melanophloia as a result of fire are relatively low. Girth growth was enhanced in wet years, particularly for larger trees (>10 cm dbh), but all trees regardless of size or woody competition levels are vulnerable to drought‐induced mortality. Overall the results suggest that variations in rainfall, especially drought‐induced mortality, have a much stronger influence on the tree demographics of E. melanophloia in a semi‐arid savanna of north‐eastern Australia than fire.     

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,...     

松针叶群数量和寿命的水平与垂直变化趋势分析

根据对巴山松针叶群体的水平和垂直变化特征及趋势进行的研究,得到如下结果：(1)单株木的针叶总数和各龄针叶总数随胸径增加而增多,随密度增加而减少;当年生针叶数量所占比例随径阶增加而增大,2年生针叶比例与此相反,而1年生和3年生针叶比例基本保持稳定。(2)单株木的径阶愈小,针叶平均寿命亦愈长;就针叶平均生存期限而言,劣势木最短,优势木次之,而平均木最长。(3)各龄级针叶群体的数量一般在树冠中部所占比例最大,下部所占比例次之,上部比例最小。在林分中处于不同地位的林木表现形式略有差异,与树冠内光照的分布有关。(4)针叶的内禀增长率随着径阶的增大、树高的增加及相对光照强度的增高而增加;树冠中针叶的平均寿命随树高增加而减小。另外,对产生上述结果的原因也进行了部分探讨。     

Morphological and phenological shoot plasticity in a Mediterranean evergreen oak facing long-term increased drought

Mediterranean trees must adjust their canopy leaf area to the unpredictable timing and severity of summer drought. The impact of increased drought on the canopy dynamics of the evergreen Quercus ilex was studied by measuring shoot growth, leaf production, litterfall, leafing phenology and leaf demography in a mature forest stand submitted to partial throughfall exclusion for 7 years. The leaf area index rapidly declined in the throughfall-exclusion plot and was 19% lower than in the control plot after 7 years of treatment. Consequently, leaf litterfall was significantly lower in the dry treatment. Such a decline in leaf area occurred through a change in branch allometry with a decreased number of ramifications produced and a reduction of the leaf area supported per unit sapwood area of the shoot (LA/SA). The leafing phenology was slightly delayed and the median leaf life span was slightly longer in the dry treatment. The canopy dynamics in both treatments were driven by water availability with a 1-year lag: leaf shedding and production were reduced following dry years; in contrast, leaf turnover was increased following wet years. The drought-induced decrease in leaf area, resulting from both plasticity in shoot development and slower leaf turnover, appeared to be a hydraulic adjustment to limit canopy transpiration and maintain leaf-specific hydraulic conductivity under drier conditions.     

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.     

Annual and spatial variation in shoot demography associated with masting in Betula grossa: comparison between mature trees and saplings

Backgrounds and Aims

Shoot demography affects the growth of the tree crown and the number of leaves on a tree. Masting may cause inter-annual and spatial variation in shoot demography of mature trees, which may in turn affect the resource budget of the tree. The aim of this study was to evaluate the effect of masting on the temporal and spatial variations in shoot demography of mature Betula grossa.

Methods

The shoot demography was analysed in the upper and lower parts of the tree crown in mature trees and saplings over 7 years. Mature trees and saplings were compared to differentiate the effect of masting from the effect of exogenous environment on shoot demography. The fate of different shoot types (reproductive, vegetative, short, long), shoot length and leaf area were investigated by monitoring and by retrospective survey using morphological markers on branches. The effects of year and branch position on demographic parameters were evaluated.

Key Results

Shoot increase rate, production of long shoots, bud mortality, length of long shoots and leaf area of a branch fluctuated periodically from year to year in mature trees over 7 years, in which two masting events occurred. Branches within a crown showed synchronized annual variation, and the extent of fluctuation was larger in the upper branches than the lower branches. Vegetative shoots varied in their bud differentiation each year and contributed to the dynamic shoot demography as much as did reproductive shoots, suggesting physiological integration in shoot demography through hormonal regulation and resource allocation.

Conclusions

Masting caused periodic annual variation in shoot demography of the mature trees and the effect was spatially variable within a tree crown. Since masting is a common phenomenon among tree species, annual variation in shoot demography and leaf area should be incorporated into resource allocation models of mature masting trees.     

Woodland recovery following drought‐induced tree mortality across an environmental stress gradient

Recent droughts and increasing temperatures have resulted in extensive tree mortality across the globe. Understanding the environmental controls on tree regeneration following these drought events will allow for better predictions of how these ecosystems may shift under a warmer, drier climate. Within the widely distributed piñon–juniper woodlands of the southwestern USA, a multiyear drought in 2002–2004 resulted in extensive adult piñon mortality and shifted adult woodland composition to a juniper‐dominated, more savannah‐type ecosystem. Here, we used pre‐ (1998–2001) and 10‐year post‐ (2014) drought stand structure data of individually mapped trees at 42 sites to assess the effects of this drought on tree regeneration across a gradient of environmental stress. We found declines in piñon juvenile densities since the multiyear drought due to limited new recruitment and high (>50%) juvenile mortality. This is in contrast to juniper juvenile densities, which increased over this time period. Across the landscape, piñon recruitment was positively associated with live adult piñon densities and soil available water capacity, likely due to their respective effects on seed and water availability. Juvenile piñon survival was strongly facilitated by certain types of nurse trees and shrubs. These nurse plants also moderated the effects of environmental stress on piñon survival: Survival of interspace piñon juveniles was positively associated with soil available water capacity, whereas survival of nursed piñon juveniles was negatively associated with perennial grass cover. Thus, nurse plants had a greater facilitative effect on survival at sites with higher soil available water capacity and perennial grass cover. Notably, mean annual climatic water deficit and elevation were not associated with piñon recruitment or survival across the landscape. Our findings reveal a clear shift in successional trajectories toward a more juniper‐dominated woodland and highlight the importance of incorporating biotic interactions and soil properties into species distribution modeling approaches.     

Seasonal leaf dynamics across a tree density gradient in a Brazilian savanna

Interactions between trees and grasses that influence leaf area index (LAI) have important consequences for savanna ecosystem processes through their controls on water, carbon, and energy fluxes as well as fire regimes. We measured LAI, of the groundlayer (herbaceous and woody plants <1-m tall) and shrub and tree layer (woody plants >1-m tall), in the Brazilian cerrado over a range of tree densities from open shrub savanna to closed woodland through the annual cycle. During the dry season, soil water potential was strongly and positively correlated with grass LAI, and less strongly with tree and shrub LAI. By the end of the dry season, LAI of grasses, groundlayer dicots and trees declined to 28, 60, and 68% of mean wet-season values, respectively. We compared the data to remotely sensed vegetation indices, finding that field measurements were more strongly correlated to the enhanced vegetation index (EVI, r ²=0.71) than to the normalized difference vegetation index (NDVI, r ²=0.49). Although the latter has been more widely used in quantifying leaf dynamics of tropical savannas, EVI appears better suited for this purpose. Our ground-based measurements demonstrate that groundlayer LAI declines with increasing tree density across sites, with savanna grasses being excluded at a tree LAI of approximately 3.3. LAI averaged 4.2 in nearby gallery (riparian) forest, so savanna grasses were absent, thereby greatly reducing fire risk and permitting survival of fire-sensitive forest tree species. Although edaphic conditions may partly explain the larger tree LAI of forests, relative to savanna, biological differences between savanna and forest tree species play an important role. Overall, forest tree species had 48% greater LAI than congeneric savanna trees under similar growing conditions. Savanna and forest species play distinct roles in the structure and dynamics of savanna–forest boundaries, contributing to the differences in fire regimes, microclimate, and nutrient cycling between savanna and forest ecosystems.     

巴山松针叶群体数量和寿命的水平与垂直变化趋势分析

根据对巴山松针叶群体的水平和垂直主特征及趋势进行的研究,得到如下结果：（１）单株木的针叶总数和各龄针总数随胸径增加而增多,随密度增加而减少,当年生针叶数量所占比例随径阶增加而增大,２年生针叶比例与此林反,而１年生和３年生针叶比例基本保持稳定。（２）单株木的径阶愈小,针叶平均寿命亦愈长;就针叶平均生存期则言,劣势本最短,成势木次之,而平均木最长。（３）各龄级针叶群体的数量一般在树冠中部所占比例最大,     

Relative Impacts of Elephant and Fire on Large Trees in a Savanna Ecosystem

Elephant and fire are considered to be among the most important agents that can modify the African savanna ecosystem. Although the synergistic relationship between these two key ecological drivers is well documented, it has proved much more difficult to establish the relative effects they have on savanna vegetation structure at a fine-scale over time. In this study, we explore the comparative impacts of fire and elephant on 2,522 individually identified large trees (≥5 m in height) in the Kruger National Park, South Africa. Data were collected from 21 transects first surveyed in April 2006 and resurveyed in November 2008, to determine the relative importance of past damage by these agents on subsequent impacts and mortality. The occurrence of fire or elephant damage in 2006 affected the amount of tree volume subsequently removed by both these agents; elephant removed more tree volume from previously burned trees and the impact of subsequent fire was higher on previously burned or elephant-utilized trees than on undamaged trees. Mortality was also affected by an interaction between previous and recent damage, as the probability of mortality was highest for trees that suffered from fire or elephant utilization after being pushed over. Subsequent fire damage, but not elephant utilization, on debarked trees also increased the probability of mortality. Mortality was twice (4.6% per annum) that of trees progressing into the ≥5 m height class, suggesting an overall decline in large tree density during the 30-month study period. The responses of large trees were species and landscape-specific in terms of sensitivity to elephant and fire impacts, as well as for levels of mortality and progression into the ≥5 m height class. These results emphasize the need for fine-scale site-specific knowledge for effective landscape level understanding of savanna dynamics.     

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.