Site‐adapted admixed tree species reduce drought susceptibility of mature European beech

Some forest‐related studies on possible effects of climate change conclude that growth potential of European beech (Fagus sylvatica L.) might be impaired by the predicted increase in future serious drought events during the growing season. Other recent research suggests that not only multiyear increment rates but also growth resistance and recovery of beech during, respectively, after dry years may differ between pure and mixed stands. Thus, we combined dendrochronological investigations and wood stable isotope measurements to further investigate the impact of neighborhood diversity on long‐term performance, short‐term drought response and soil water availability of European beech in three major geographic regions of Germany. During the last four decades, target trees whose competitive neighborhood consisted of co‐occurring species exhibited a superior growth performance compared to beeches in pure stands of the same investigation area. This general pattern was also found in exceptional dry years. Although the summer droughts of 1976 and 2003 predominantly caused stronger relative growth declines if target trees were exposed to interspecific competition, with few exceptions they still formed wider annual rings than beeches growing in close‐by monocultures. Within the same study region, recovery of standardized beech target tree radial growth was consistently slower in monospecific stands than in the neighborhood of other competitor species. These findings suggest an improved water availability of beech in mixtures what is in line with the results of the stable isotope analysis. Apparently, the magnitude of competitive complementarity determines the growth response of target beech trees in mixtures. Our investigation strongly suggest that the sensitivity of European beech to environmental constrains depends on neighborhood identity. Therefore, the systematic formation of mixed stands tends to be an appropriate silvicultural measure to mitigate the effects of global warming and droughts on growth patterns of Fagus sylvatica.     

Patterns of drought tolerance in major European temperate forest trees: climatic drivers and levels of variability

The future performance of native tree species under climate change conditions is frequently discussed, since increasingly severe and more frequent drought events are expected to become a major risk for forest ecosystems. To improve our understanding of the drought tolerance of the three common European temperate forest tree species Norway spruce, silver fir and common beech, we tested the influence of climate and tree‐specific traits on the inter and intrasite variability in drought responses of these species. Basal area increment data from a large tree‐ring network in Southern Germany and Alpine Austria along a climatic cline from warm‐dry to cool‐wet conditions were used to calculate indices of tolerance to drought events and their variability at the level of individual trees and populations. General patterns of tolerance indicated a high vulnerability of Norway spruce in comparison to fir and beech and a strong influence of bioclimatic conditions on drought response for all species. On the level of individual trees, low‐growth rates prior to drought events, high competitive status and low age favored resilience in growth response to drought. Consequently, drought events led to heterogeneous and variable response patterns in forests stands. These findings may support the idea of deliberately using spontaneous selection and adaption effects as a passive strategy of forest management under climate change conditions, especially a strong directional selection for more tolerant individuals when frequency and intensity of summer droughts will increase in the course of global climate change.     

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.     

Anthropogenic‐driven rapid shifts in tree distribution lead to increased dominance of broadleaf species

Over the past century, major shifts in the geographic distribution of tree species have occurred in response to changes in land use and climate. We analyse species distribution and abundance from about 33 000 forest inventory plots in Spain sampled twice over a period of 10–12 years. We show a dominance of range contraction (extinction), and demographic decline over range expansion (colonization), with seven of 11 species exhibiting extinction downhill of their distribution. Contrary to expectations, these dynamics are not always consistent with climate warming over the study period, but result from legacies in forest structure due to past land use change and fire occurrence. We find that these changes have led to the expansion of broadleaf species (i.e. family Fagaceae) over areas formerly dominated by conifer species (i.e. family Pinaceae), due to the greater capacity of the former to respond to most disturbances and their higher competitive ability. This recent and rapid transition from conifers to broadleaves has important implications in forest dynamics and ecosystem services they provide. The finding raises the question as to whether the increasing dominance of relatively drought‐sensitive broadleaf species will diminish resilience of Mediterranean forests to very likely drier conditions in the future.     

Global phylogeography of the genus Capreolus (Artiodactyla: Cervidae), a Palaearctic meso‐mammal

Areas of sympatry and hybridization of closely related species can be difficult to assess through morphological differences alone. Species which coexist and are similar morphologically may be distinguished only with molecular techniques. The roe deer (Capreolus spp.) is a meso‐mammal having a Palaearctic distribution, with two closely related species: the European C. capreolus and the Siberian C. pygargus. We analysed mtDNA sequences from 245 individuals, sampled through all the entire range of the genus, to investigate the distribution of genetic lineages and outline phylogeographical patterns. We found that: (1) a C. pygargus lineage occurs in Poland and Lithuania, much farther west than the area which so far was believed its westernmost limit; (2) no haplotype of this C. pygargus lineage matches any found in East Europe and Asia – this should rule out human introductions and may indicate Pleistocene–Holocene migrations from the east; (3) no geographical structuring of C. pygargus lineages occurs, questioning the existence of putative subspecies; (4) several genetic lineages of C. capreolus can be recognized, consistent with the existence of two subspecies, respectively in central–southern Italy and southern Spain. Coalescence times suggest that intraspecific variation in C. capreolus and C. pygargus developed approximately 100–10 kya. The extant mitochondrial lineages pre‐dated the Last Glacial Maximum. Capreolus pygargus must have moved westward to Central Europe, where at least one genetic lineage still survives, coexisting with C. capreolus. © 2013 The Linnean Society of London     

鼎湖山针阔叶混交林4种优势树种树干液流特征

运用Granier热消散式探针法,对鼎湖山自然保护区针阔叶混交林4种优势树种(马尾松、木荷、锥栗和广东润楠)的树干液流密度进行了长期连续观测,并同步监测林分的环境因子(光合有效辐射(PAR)、空气温度(T)、空气湿度(RH)、土壤含水量)。结果表明:(1)4种优势树种的边材面积(As)与胸径(DBH)均存在显著相关关系(P0.05);(2)各优势树种树干液流均呈现"昼高夜低"单峰曲线,且液流速率存在明显的季节性差异;(3)无论湿季还是干季,光合有效辐射(PAR)和水汽压亏缺(VPD)均为控制蒸腾的主要驱动因子;(4)4种优势树种湿季平均日蒸腾量高于干季,马尾松、木荷、锥栗、广东润楠湿季平均日蒸腾量分别为29.52、39.29、30.40、9.41 kg H2O/d,干季分别为20.91、24.84、24.26、8.43 kg H2O/d,干季和湿季的平均日蒸腾量(kg H2O/d)大小均为木荷锥栗马尾松广东润楠,这种种间差异是由边材面积大小和树种本身的生物学特性共同决定的。     

Acclimation of photosynthetic capacity to irradiance in tree canopies in relation to leaf nitrogen concentration and leaf mass per unit area

The observation of acclimation in leaf photosynthetic capacity to differences in growth irradiance has been widely used as support for a hypothesis that enables a simplification of some soil‐vegetation‐atmosphere transfer (SVAT) photosynthesis models. The acclimation hypothesis requires that relative leaf nitrogen concentration declines with relative irradiance from the top of a canopy to the bottom, in 1 : 1 proportion. In combination with a light transmission model it enables a simple estimate of the vertical profile in leaf nitrogen concentration (which is assumed to determine maximum carboxylation capacity), and in combination with estimates of the fraction of absorbed radiation it also leads to simple ‘big‐leaf’ analytical solutions for canopy photosynthesis. We tested how forests deviate from this condition in five tree canopies, including four broadleaf stands, and one needle‐leaf stand: a mixed‐species tropical rain forest, oak (Quercus petraea (Matt.) Liebl), birch (Betula pendula Roth), beech (Fagus sylvatica L.) and Sitka spruce (Picea sitchensis (Bong.) Carr). Each canopy was studied when fully developed (mid‐to‐late summer for temperate stands). Irradiance (Q, µmol m^?2 s^?1) was measured for 20 d using quantum sensors placed throughout the vertical canopy profile. Measurements were made to obtain parameters from leaves adjacent to the radiation sensors: maximum carboxylation and electron transfer capacity (V_a, J_a, µmol m^?2 s^?1), day respiration (R_da, µmol m^?2 s^?1), leaf nitrogen concentration (N_m, mg g^?1) and leaf mass per unit area (L_a, g m^?2). Relative to upper‐canopy values, V_a declined linearly in 1 : 1 proportion with N_a. Relative V_a also declined linearly with relative Q, but with a significant intercept at zero irradiance (P < 0·01). This intercept was strongly related to L_a of the lowest leaves in each canopy (P < 0·01, r² = 0·98, n= 5). For each canopy, daily lnQ was also linearly related with lnV_a(P < 0·05), and the intercept was correlated with the value for photosynthetic capacity per unit nitrogen (PUN: V_a/N_a, µmol g^?1 s^?1) of the lowest leaves in each canopy (P < 0·05). V_a was linearly related with L_a and N_a(P < 0·01), but the slope of the V_a : N_a relationship varied widely among sites. Hence, whilst there was a unique V_a : N_a ratio in each stand, acclimation in V_a to Q varied predictably with L_a of the lowest leaves in each canopy. The specific leaf area, L_m(cm² g^?1), of the canopy‐bottom foliage was also found to predict carboxylation capacity (expressed on a mass basis; V_m, µmol g^?1 s^?1) at all sites (P < 0·01). These results invalidate the hypothesis of full acclimation to irradiance, but suggest that L_a and L_m of the most light‐limited leaves in a canopy are widely applicable indicators of the distribution of photosynthetic capacity with height in forests.     

Forest structure and tree species distribution in relation to topography-mediated heterogeneity of soil nitrogen and light at the forest floor

We investigated the patterns of soil nitrogen (N) and forest floor light availability, forest structure and tree species distribution along a topographic gradient on a 200-m mountain slope in a cool-temperate deciduous broad-leaved forest in Japan. Rates of soil N mineralization and nitrification decreased from lower to upper slope positions, revealing that N availability decreased up the slope. Maximum tree height and above-ground biomass were greater on the lower than the upper parts of the slope. Canopy openness, an index of light availability at the forest floor, increased up the slope. Tree species could be placed into three groups according to their distribution patterns on the slope. Ridge and valley species were distributed on the upper and lower parts of the slope, respectively, whereas uniform species were distributed over the entire slope. Topography-mediated resource gradients of soil N and light may be important determinants of species distribution patterns and forest regeneration, and the results of this study imply that the determinants of the regeneration process differ between the lower and upper parts of a slope. The former may be relatively light limited and the latter may be soil N limited. Valley species may have a greater ability to compete for light, whereas ridge species have a greater ability to compete for soil N. The broad distribution of uniform species probably reflects an ability to effectively compete for both light and soil N.     

Environmental effects on fine‐scale spatial genetic structure in four Alpine keystone forest tree species

Genetic responses to environmental changes take place at different spatial scales. While the effect of environment on the distribution of species' genetic diversity at large geographical scales has been the focus of several recent studies, its potential effects on genetic structure at local scales are understudied. Environmental effects on fine‐scale spatial genetic structure (FSGS) were investigated in four Alpine conifer species (five to eight populations per species) from the eastern Italian Alps. Significant FSGS was found for 11 of 25 populations. Interestingly, we found no significant differences in FSGS across species but great variation among populations within species, highlighting the importance of local environmental factors. Interannual variability in spring temperature had a small but significant effect on FSGS of Larix decidua, probably related to species‐specific life history traits. For Abies alba, Picea abies and Pinus cembra, linear models identified spring precipitation as a potentially relevant climate factor associated with differences in FSGS across populations; however, models had low explanatory power and were strongly influenced by a P. cembra outlier population from a very dry site. Overall, the direction of the identified effects is according to expectations, with drier and more variable environments increasing FSGS. Underlying mechanisms may include climate‐related changes in the variance of reproductive success and/or environmental selection of specific families. This study provides new insights on potential changes in local genetic structure of four Alpine conifers in the face of environmental changes, suggesting that new climates, through altering FSGS, may also have relevant impacts on plant microevolution.     

Long‐term effects of climate change on carbon storage and tree species composition in a dry deciduous forest

Forest vegetation and soils have been suggested as potentially important sinks for carbon (C) with appropriate management and thus are implicated as effective tools in stabilizing climate even with increasing anthropogenic release of CO₂. Drought, however, which is often predicted to increase in models of future climate change, may limit net primary productio (NPP) of dry forest types, with unknown effects on soil C storage. We studied C dynamics of a deciduous temperate forest of Hungary that has been subject to significant decreases in precipitation and increases in temperature in recent decades. We resampled plots that were established in 1972 and repeated the full C inventory by analyzing more than 4 decades of data on the number of living trees, biomass of trees and shrubs, and soil C content. Our analyses show that the decline in number and biomass of oaks started around the end of the 1970s with a 71% reduction in the number of sessile oak stems by 2014. Projected growth in this forest, based on the yield table's data for Hungary, was 4.6 kg C/m². Although new species emerged, this new growth and small increases in oak biomass resulted in only 1.9 kg C/m² increase over 41 years. The death of oaks increased inputs of coarse woody debris to the surface of the soil, much of which is still identifiable, and caused an increase of 15.5%, or 2.6 kg C/m², in the top 1 m of soil. Stability of this fresh organic matter input to surface soil is unknown, but is likely to be low based on the results of a colocated woody litter decomposition study. The effects of a warmer and drier climate on the C balance of forests in this region will be felt for decades to come as woody litter inputs decay, and forest growth remains impeded.     

Continental‐scale tree‐ring‐based projection of Douglas‐fir growth: Testing the limits of space‐for‐time substitution

A central challenge in global change research is the projection of the future behavior of a system based upon past observations. Tree‐ring data have been used increasingly over the last decade to project tree growth and forest ecosystem vulnerability under future climate conditions. But how can the response of tree growth to past climate variation predict the future, when the future does not look like the past? Space‐for‐time substitution (SFTS) is one way to overcome the problem of extrapolation: the response at a given location in a warmer future is assumed to follow the response at a warmer location today. Here we evaluated an SFTS approach to projecting future growth of Douglas‐fir (Pseudotsuga menziesii), a species that occupies an exceptionally large environmental space in North America. We fit a hierarchical mixed‐effects model to capture ring‐width variability in response to spatial and temporal variation in climate. We found opposing gradients for productivity and climate sensitivity with highest growth rates and weakest response to interannual climate variation in the mesic coastal part of Douglas‐fir's range; narrower rings and stronger climate sensitivity occurred across the semi‐arid interior. Ring‐width response to spatial versus temporal temperature variation was opposite in sign, suggesting that spatial variation in productivity, caused by local adaptation and other slow processes, cannot be used to anticipate changes in productivity caused by rapid climate change. We thus substituted only climate sensitivities when projecting future tree growth. Growth declines were projected across much of Douglas‐fir's distribution, with largest relative decreases in the semiarid U.S. Interior West and smallest in the mesic Pacific Northwest. We further highlight the strengths of mixed‐effects modeling for reviving a conceptual cornerstone of dendroecology, Cook's 1987 aggregate growth model, and the great potential to use tree‐ring networks and results as a calibration target for next‐generation vegetation models.     

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.     

Red oak seedlings as indicators of deer browse pressure: Gauging the outcome of different white‐tailed deer management approaches

After decades of high deer populations, North American forests have lost much of their previous biodiversity. Any landscape‐level recovery requires substantial reductions in deer herds, but modern societies and wildlife management agencies appear unable to devise appropriate solutions to this chronic ecological and human health crisis. We evaluated the effectiveness of fertility control and hunting in reducing deer impacts at Cornell University. We estimated spring deer populations and planted Quercus rubra seedlings to assess deer browse pressure, rodent attack, and other factors compromising seedling performance. Oak seedlings protected in cages grew well, but deer annually browsed ≥60% of unprotected seedlings. Despite female sterilization rates of >90%, the deer population remained stable. Neither sterilization nor recreational hunting reduced deer browse rates and neither appears able to achieve reductions in deer populations or their impacts. We eliminated deer sterilization and recreational hunting in a core management area in favor of allowing volunteer archers to shoot deer over bait, including at night. This resulted in a substantial reduction in the deer population and a linear decline in browse rates as a function of spring deer abundance. Public trust stewardship of North American landscapes will require a fundamental overhaul in deer management to provide for a brighter future, and oak seedlings may be a promising metric to assess success. These changes will require intense public debate and may require new approaches such as regulated commercial hunting, natural dispersal, or intentional release of important deer predators (e.g., wolves and mountain lions). Such drastic changes in deer management will be highly controversial, and at present, likely difficult to implement in North America. However, the future of our forest ecosystems and their associated biodiversity will depend on evidence to guide change in landscape management and stewardship.     

不同氮添加量和添加方式对南亚热带4个主要树种幼苗生长的影响

为阐明南亚热带4个主要树种——海南红豆(Ormosia pinnata)、马占相思(Acacia mangium)、木荷(Schima superba)和马尾松(Pinus massoniana)幼苗生长对不同氮添加量和添加方式的响应差异, 进行了幼苗模拟氮添加实验。实验设置3个氮添加水平(对照: 背景大气氮沉降量5.6 g N·m^-2·a^-1, 中氮: 15.6 g N·m^-2·a^-1, 高氮: 20.6 g N·m^-2·a^-1), 每个水平分两种添加方式(幼苗冠层施氮和土壤表层施氮), 共6个处理: (1)土壤对照(S-CK); (2)土壤中氮(S-MN); (3)土壤高氮(S-HN); (4)冠层对照(C-CK); (5)冠层中氮(C-MN); (6)冠层高氮(C-HN), 每个处理设置6个重复。研究结果表明: 不同氮添加量下, 土壤施氮和冠层施氮对植物幼苗生长的影响不同, 氮添加量、氮添加方式和物种3个因子之间存在显著的交互效应。与对照相比, S-MN增加了马占相思和木荷幼苗的生物量, 降低了马尾松的株高和生物量, 而C-MN仅增加了马占相思的生物量, 对其他3个树种没有影响; S-HN增加了马占相思的生物量, 显著降低了马尾松的基径、株高和生物量(p < 0.01), C-HN增加了马占相思、木荷和马尾松的基径、株高和生物量(p < 0.01)。不同氮添加量和氮添加方式对幼苗生长的影响因物种而异, 所有氮处理下海南红豆和马占相思的生长均明显快于木荷和马尾松; 木荷和马尾松幼苗的生长在两种氮添加方式间差异显著, 冠层施氮比土壤施氮对其幼苗生长的促进作用更大。由此可见: 在氮沉降背景下, 阔叶豆科植物(海南红豆、马占相思)比阔叶非豆科植物(木荷)生长快; 阔叶树种(海南红豆、马占相思和木荷)比针叶树种(马尾松)生长快。在长期氮沉降环境下, 不同物种生长的差异响应有可能导致亚热带森林物种组成发生变化。