To what extent is altitudinal variation of functional traits driven by genetic adaptation in European oak and beech?

The phenotypic responses of functional traits in natural populations are driven by genetic diversity and phenotypic plasticity. These two mechanisms enable trees to cope with rapid climate change. We studied two European temperate tree species (sessile oak and European beech), focusing on (i) in situ variations of leaf functional traits (morphological and physiological) along two altitudinal gradients and (ii) the extent to which these variations were under environmental and/or genetic control using a common garden experiment. For all traits, altitudinal trends tended to be highly consistent between species and transects. For both species, leaf mass per area displayed a positive linear correlation with altitude, whereas leaf size was negatively correlated with altitude. We also observed a significant increase in leaf physiological performance with increasing altitude: populations at high altitudes had higher maximum rates of assimilation, stomatal conductance and leaf nitrogen content than those at low altitudes. In the common garden experiment, genetic differentiation between populations accounted for 0-28% of total phenotypic variation. However, only two traits (leaf mass per area and nitrogen content) exhibited a significant cline. The combination of in situ and common garden experiments used here made it possible to demonstrate, for both species, a weaker effect of genetic variation than of variations in natural conditions, suggesting a strong effect of the environment on leaf functional traits. Finally, we demonstrated that intrapopulation variability was systematically higher than interpopulation variability, whatever the functional trait considered, indicating a high potential capacity to adapt to climate change.     

Genetic and functional leaf traits variability of Quercus laurina along an oak diversity gradient in Mexico

The ecological literature has documented the effects of plant hybridization on phenotypic variation, and dominant, intermediate, or novel morphological, chemical and physiological traits in hybrids. It is important to understand the ecological consequences of hybridization by evaluating their impact on phenotypic expression of functional traits. We evaluated the relationship between genetic diversity of Quercus laurina and functional foliar traits along an oak diversity gradient. We selected five study sites that represent an oak diversity gradient where Q. laurina is present. Using chloroplast and nuclear microsatellites, we evaluated genetic diversity, measured functional foliar traits of Q. laurina in each site and assessed the effects of local climate variables on the oak community and functional traits. We found a greater abundance of Q. laurina in all study sites. We did not find a relationship between the number of accompanying red oak species and the population genetic diversity in Q. laurina, but higher genetic diversity was found in all study sites in comparison with European oak species. Sites with more oak species had more variation of foliar functional traits. Our results do not support the hypothesis that predicts higher levels of genetic diversity of Q. laurina in communities with greater oak diversity from the same section, but we demonstrated an increase in the foliar functional traits of Q. laurina associated with oak richness and climate variables. We highlight the need to consider environmental and ecological variables linkages as regulatory mechanisms of the phenotypic plasticity expressed in changes of some functional attributes of oaks.

     

Pinus chiapensis, a keystone species: Genetics,ecology, and conservation

We present an overview of recent studies carried out on Pinus chiapensis (Mart.) Andresen (Pinus strobus var. chiapensis Mart.) and provide management and conservation recommendations. Because of its wood quality, and being commonly used by 12 ethnic groups, this pine is an outstanding forest resource at mid-altitude humid mountains of southern Mexico and Guatemala. P. chiapensis appears to be a distinctive species, closely related with North American white pines, and a potential valuable resource for establishing breeding programs with such species. P. chiapensis is the most abundant tree species in early successional stands of the tropical montane cloud forest playing a key role in ecosystem regeneration particularly in areas managed under slash-and-burn practices. However, many natural stands of this pine are severely reduced. Molecular studies based on isozymes and DNA markers reveal low genetic diversity, the lowest compared with its closest relatives (P. ayacahuite, P. monticola and P. strobus). Heterozygosity and seed viability increase significantly with population size, and inbreeding depression appears to significantly decrease seed viability, suggesting the involvement of genetic factors on population decline. Low population size is associated with both lack of perturbation in well-preserved habitats and high deforestation rates in severely disturbed habitats. Conservation and management practices require preserving and restoring connections between suitable habitats to enhance gene flow between populations, and on careful programs that monitor and control slash-and-burn practices. Restoration practices should use seeds from as many tree sources as possible to reduce inbreeding risks. Spline climate models predict significant increases in temperature, decreases in precipitation and consequently an increase of aridity along the range of P. chiapensis. Thus, assisted migration would be needed to match present genotypes to forecasted climate changes.     

林芝云杉天然群体针叶与种实的变异及其地理趋势

在西藏地区11个有代表性的林芝云杉天然群体内分别随机抽取15～30个个体为试材,测量了针叶与种实共9个表型性状,采用统计分析方法研究了在群体内和群体间的变异和分化趋势.结果显示:林芝云杉9个表型性状在群体内和群体间的差异显著.群体间表型分化系数均值为29.55％,与云杉属其他树种相比处于中等水平;群体内的方差分量占70...     

Nucleotide polymorphisms associated with climate,phenology and physiological traits in European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.)

Although European beech (Fagus sylvatica L.) is one of the most widespread and ecologically and commercially most important deciduous trees in Europe, little is known about its adaptive genetic variation. We explored single-nucleotide polymorphism (SNP) variation in candidate genes for budburst and drought resistance in beech populations sampled across most of the distribution range, represented in an international provenance trial. SNP variation was monitored for six candidate genes, in 114 individuals from 19 natural populations. Population structure was deduced from the analysis of 20 nuclear microsatellite markers. Different methods to detect imprints of natural selection were used (F _ST-outlier, SNP-climate regression, association tests). The F _ST-outlier approach identified the COV gene with unambiguous signal of selection, which is an orthologue of Arabidopsis gene for a membrane protein previously reported as phenology-related. Based on environmental association analysis at the population level, the dehydrin gene was found associated with drought-related climatic variables. At the individual level, dehydrin gene also showed a significant association with chlorophyll fluorescence parameters, which are considered stress markers. The importance of the knowledge of physiological variation and geographical patterns of adaptive genetic variation for guiding reproductive materials transfer under climate change is stressed.     

Effects of treefall gaps created by windthrow on bat assemblages in a temperate forest

Determining the way in which spatial distribution and diversity of forest-dwelling mammals varies with natural disturbance is essential to understanding the spatial dynamics of mammal assemblages in forests. Bats are the only forest-dwelling mammals capable of true flight. At a local scale, bat flight ability, which may be related to ecomorphological traits, is an important factor influencing spatial distribution. We tested two postulates: (1) the spatial distribution of bats is affected by sizes of forest gaps created by natural disturbances and (2) species-specific responses can be predicted from bat ecomorphological traits (aspect ratio (AR) and wingtip shape index (WT)) that influence bat flight ability. We found that sizes of forest gaps affected the occurrence of each bat species and species richness of bats at local scales; species-specific responses were related to the ecomorphological traits of individual species. Bat species with high AR and low WT were not affected by variation in canopy gap size. In contrast, bat species with low AR and high WT responded negatively to gap size, and those with intermediate AR and WT responded positively to canopy gap size at sites with small-sized gaps but responded negatively to large-sized gaps. Overall bat species richness responded negatively to gap size. Thus, ecomorphological traits may be important determinants of bat spatial distributions and species diversity at local scales in disturbed habitats. In this study, forest edges might have been undersampled due to the location of bat detectors. However, this potential undersampling should not have affected the interpretation of occurrence patterns of bat species responding to gap size. Our results imply that bat conservation efforts in forest lands should take into consideration specific responses related to ecomorphological traits of species inhabiting an area. The results also suggest that quantifying the effects of natural disturbances on bat assemblages may provide a knowledge base for forest management to minimize the impacts of unavoidable anthropogenic disturbances on bat species diversity. Rare or infrequent natural disturbances can provide models for forest management aimed at maintaining bat species diversity.     

Morphological and genetic differentiation in isolated populations of Mexican beech Fagus grandifolia subsp. mexicana

Mexican beech [Fagus grandifolia subsp.mexicana(Martinez) A.E.Murray] is a subspecies endemic to the Sierra Madre Oriental Mountains and considered endangered due to the low density of its populations and high degree of habitat fragmentation and environmental specificity.Because its morphological and genetic variation is associated with its ability to adapt to changes in environmental conditions,the objective of this study was to determine whether phenotypic and genotypic variation exist,and it relationships with population reduction events.In four beech populations in the states of Hidalgo and Veracruz,we analyzed 11 morphological variables for leaves and 6 micros atellite markers.The morphological variables that to discriminate between populations were related to the size of the leaf,but a robust differentiation pattern was not found,given that independent groups of leaves were identified.The populations located closest to each other,had greater genetic variation and less genetic distance;populations in the extreme north and south had the lowest genetic variation.Genetic differentiation among populations was associated with reduction in population size.In the 3 localities in Hidalgo,recent bottlenecks were identified,and in Veracruz,an old bottleneck was found.Variation in leaf morphology and genetic structure of Mexican beech populations could be the result of a combination of various geographical,climate and ecological factors.     

Plant functional types rather than climate or soil determine leaf traits in the forest biomes of eastern China

Nitrogen (N) has great ecological importance, but the biogeographic pattern across forest biomes in China has only recently been explored. Here we conducted a systematic census of leaf C and N following the same protocol to explore the variations of leaf traits, and their possible responses to plant functional types (PFTs) and environmental factors. Results showed that leaf traits varied substantially across biomes, and the relationships of PFTs to climatic factors were stronger than those of PFTs versus soil nutrient proxies, indicating that plant species composition might be a better predictor of plant species distribution with climate than leaf traits. Soil nutrient proxies explained more variation of leaf traits than climate, which demonstrates that leaf traits reflect important aspects of plant responses to soil nutrients. Importantly, partial general linear models analyses found that PFTs showed the greatest direct influence for leaf traits, and climate and soil affected leaf traits mainly through the change in plant species composition rather than having direct impacts. Hence, we concluded that leaf traits were largely controlled by PFTs rather than climate or soil at the biome scale. The results favored the species composition hypothesis, indicating that leaf nutrient concentration is mainly determined by PFTs.     

Effect of patch size and isolation on mating patterns and seed production in an urban population of Chinese pine (Pinus tabulaeformis Carr.)

Fragmentation is a critical issue for tree populations because the creation of small patches can reduce local population size and increase isolation, both of which can promote inbreeding and its negative consequences, as well as loss of genetic diversity. To test the hypothesis that patch size and isolation influence mating patterns or seed production in forest trees, we utilize the spatial array of trees of an urban population of Chinese Pine (Pinus tabulaeformis Carr.) planted in patches around Beijing. Our design includes 28 urban patches, with patch size ranging from 1 to 2000 adult trees and isolation (edge distance index) ranging from 37.5 m to 245.8 m. We examined the average number of seeds per cone and percentage of viable seed per cone for each patch as measures of seed production. By utilizing seven paternally inherited chloroplast microsatellite loci, we estimated the mating pattern parameters for each patch, including the level of selfing, the amount of immigrant pollen and the effective number of pollen sources (N_ep). Using a general linear model selection procedure based on AIC value, we found patch size was the best predictor of the selfing and immigration rate; smaller patches had a higher selfing and immigration rate. Small patches with one adult had relatively high N_ep which indicates connectivity among urban patches. However, due to the reduced amount of immigrant pollen and limited diversity of local pollen, intermediate sized patches (with 5–10 adults) had the lowest N_ep among the study patches. For patches with more than 10 adults, N_ep was increased with patch size. The percentage of viable seeds per cone significantly decreased with patch size, indicating a possible negative consequence of inbreeding. The effect of patch size on mating patterns and seed production suggests that the patches of trees experience less connectivity than trees within continuous forest. These findings indicate that forest management practices should emphasize the maintenance of an optimal patch size because, despite the fact that tree species show the potential for long distance pollen movement, the number of local trees strongly influences the mating patterns.     

秦岭西段锐齿栎群落林木个体大小分布特征及物种多样性

对秦岭西段小陇山林区腹地和林缘区的锐齿栎群落的大小分布和多样性进行了分析,结果表明:林区腹地分布区锐齿栎群落的个体较多,锐齿栎成年林木树高频度分布的峰值出现在7 - 19 m间,占总数的73.54%, 3 m以下的幼苗幼树占总数的18.43%,锐齿栎幼苗的萌发受中成林木、灌木的影响较大。在林缘分布区内,小于3 m以下的幼苗幼树与林区腹地分布相类似,但3 m以上各树高阶的数量都很少,特别是成熟林木较少,群落处于不稳定状态。林区腹地锐齿栎群落不同层次的物种组成丰富,物种数均高于林缘区,林区腹地锐齿栎群落多样性灌木层>草本层>乔木层,林缘区锐齿栎群落多样性草本层>灌木层>乔木层。     

Canopy gaps and regeneration in old-growth Oriental beech (Fagus orientalis Lipsky) stands, northern Iran

Virgin beech Fagus orientalis forests in northern Iran provide a unique opportunity to study the disturbance regimes of forest ecosystems without human influence. The aim of this research was to describe characteristics of natural canopy gaps and gap area fraction as an environmental influence on the success of beech seedling establishment in mature beech stands. All canopy gaps and related forest parameters were measured within three 25 ha areas within the Gorazbon compartment of the University of Tehran’s Kheyrud Experimental Forest. An average of 3 gaps/ha occurred in the forest and gap sizes ranged from 19 to 1250 m² in size. The most frequent (58%) canopy gaps were <200 m². In total, canopy gaps covered 9.3% of the forest area. Gaps <400 m² in size were irregular in shape, but larger gaps did not differ significantly in shape from a circle. Most gaps (41%) were formed by a single tree-fall event and beech made up 63% of gap makers and 93% of gap fillers. Frequency and diversity of tree seedlings were not significantly correlated with gap size. The minimum gap size that contained at least one beech gap-filling sapling (<1.3 m tall) was 23.7 m². The median gap size containing at least one beech gap-filling sapling was 206 m² and the maximum size was 1808 m². The management implications from our study suggest that the creation of small and medium sized gaps in mixed beech forest should mimic natural disturbance regimes and provide suitable conditions for successful beech regeneration.     

南北样带温带区栎属树种幼苗功能性状的变异研究

植物功能性状由生境条件和遗传因素共同决定,田间控制实验能够揭示植物功能性状在种内和种间的变异程度.采集南北样带温带区8个地点6种栎属树种的种子,通过田间栽培试验,以均一环境下1年生幼苗为对象,研究比较种内及种间比叶重、单位叶面积光合速率、单位叶重量光合速率、总干物质含量和根冠比5种功能性状的变化差异,并且分析幼苗功能性状与种子性状的关系.结果发现:栎属树种种间幼苗的功能性状变异系数高于种内,并且种内功能性状的差异较小,种间差异性较大,而比叶重和叶片光合速率无论在种间还是种内,其变异率都较小;总干物质量和根冠比与种子质量显著相关,随种子质量增加幼苗的总干物质量和根冠比呈现增加的趋势,比叶重和叶片光合速率与种子质量的相关性较小.     

Range-wide conservation of <Emphasis Type="Italic">Pinus aristata</Emphasis>: a genetic collection with ecological context for proactive management today and resources for tomorrow

Tree species are highly vulnerable to anthropogenic environmental change, and are increasingly being challenged by non-native pests and climate change. Rocky Mountain bristlecone pines are long-lived, exhibit delayed maturation, have low genetic diversity, and inhabit cold, high-elevation environments. They are threatened by the non-native disease white pine blister rust, warming temperatures, changing precipitation patterns, and altered disturbance regimes. The goal of this work was to (1) sample the range-wide genetic diversity of bristlecone pine for ex situ seed and tissue collections and research before the populations are impacted, (2) assess the health and ecological conditions of the species across its geographic range (61 stands) to provide a baseline by which to evaluate future changes, and (3) assess relationships between stand and regeneration characteristics and topographic, geographic, and climatic factors to identify vulnerabilities and proactive management interventions to increase population resilience. Local variation in topoclimate was strongly related to stand structure, composition, health, and regeneration. Bristlecone pine currently showed only minor impacts from insects and pathogens, but relationships between cone production and regeneration with climate variables suggest vulnerabilities to projected climate change. Both cone production and seedling regeneration were also linked to stand structure and ground cover that provide opportunities for proactive in situ management to increase population size to mitigate potential future climate- and pathogen-driven declines. These efforts represent the first proactive coordinated range-wide genetic collection design and forest health assessment for a threatened, but not yet impacted, tree species and offers a model for other species at risk.     

Seasonal and interannual ecophysiological responses of beech (Fagus sylvatica) at its south-eastern distribution limit in Europe

Due to its wide European distribution and its drought-susceptibility, beech (Fagus sylvatica L.) received intensive attention recently in the light of global warming. Contrary to central European beech ecosystems, little is known about the ecophysiology of beech at its south-eastern European distribution limit. Here we tested whether climatic fluctuations during a three-year period affected the ecophysiology of a beech site in Greece. Attention was paid at comparing our findings to the intense effects the 2003 extreme drought had on beech forests in central Europe.     

Present and forecasted xeric climatic limits of beech and sessile oak distribution at low altitudes in Central Europe

• Introduction   

Xeric (trailing) forest range limits are particularly vulnerable to impacts of predicted climate change. Regional modelling studies contribute to the identification of potential local climatic threats and may support appropriate management strategies.     

Contemporary climate influence on variability patterns of Anadenanthera colubrina var.cebil,a key species in seasonally dry tropical forests

The distribution of many plant species has been shaped by climate changes,and their current phenotypic and genetic variability reflect microclimatically suitable habi-tats.This study relates contemporary climate to variability patterns of phenotypic traits and molecular markers in the Argentinean distribution of Anadenanthera colubrina var.cebil,as well as to identify the most relevant phenotypic trait or molecular marker associated with those patterns.Individuals from four populations in both biogeographic provinces,Paranaense and Yungas,were investigated.Multivariate analyses and multiple linear regressions were carried out to determine relationships among phenotypic traits and nuclear microsatellites,respectively,to climatic variables,and to identify the phenotypic traits as well as nuclear microsatellite loci most sensitive to climate.Two and three clusters of individuals were detected based on genetic and phenotypic data,respectively.Only clusters based on genetic data reflected the biogeographic origin of individuals.Reproductive traits were the most relevant indi-cators of climatic effects.One microsatellite locus Ac41.1 appeared to be non-neutral presenting a strong correlation with climate variable temperature seasonality.Our findings show complex patterns of genetic and phenotypic variability in the Argentinean distribution of A.colubrina var.cebil related to the present or contemporary climate,and provides an example for an integrative approach to better understand climate impact on contemporary genetic and phenotypic variability in light of global climate change.     

Regional impact assessment on forest structure and functions under climate change—the Brandenburg case study

A forest simulation model has been applied in a regional impact assessment to investigate impacts of climate change on forest structure and function in the Federal state of Brandenburg, Germany. The forest model FORSKA-M was linked to a GIS that included soil, groundwater table and land-use maps. Two climate scenarios (current climate and a climate change of 1.5 K temperature increase which is combined with a precipitation decrease of 10–20% on average) for 40 meteorological stations in and around Brandenburg were used to assess the sensitivity of species composition to climate change. Furthermore, the implications of vegetation changes for other forest functions were analysed by means of several indicators. To evaluate the impacts of climate change on biodiversity, measures of species diversity (Shannon’s and Simpson’s index) and habitat and structural diversity (Seibert’s index) were applied. The evaluation of impacts on groundwater recharge of natural and managed forests was carried out using the soil water balance model of FORSKA-M.At first, model simulations of the potential natural vegetation (PNV) on the whole area of Brandenburg with different climate scenarios were analysed. The results indicated that climatic warming would lead to a shift in the natural species composition in Brandenburg towards more drought tolerant species. The simulated diversity of the forests would be reduced, and groundwater recharge would be decreased.The majority of forests in the state of Brandenburg have been managed intensively in the past. At present, large areas of Brandenburg’s forests are dominated by pure stands of Scots pine, but current forest management practice aims at increasing the share of deciduous and mixed forests. In order to analyse the possible consequences of climate change on forest management, forest inventory data were used to initialise FORSKA-M with representative forest stands. Simulation experiments with three different management scenarios showed that the short to mid-term effects of climatic change in terms of species composition were not as severe as expected. However, the comparison of different diversity measures indicates a decrease in the species diversity in contrast to an increase in habitat diversity under climate warming. Furthermore, a decrease in productivity and groundwater recharge was simulated under the climate change scenario.The regional impact assessment corroborated the high sensitivity of natural forests in the region to the projected climatic change and it underlined the importance of adaptive management strategies to help forestry to cope with climatic change.     

Drought matters - Declining precipitation influences growth of Fagus sylvatica L. and Quercus robur L. in north-eastern Germany

For north-eastern Germany regional climate models project rising temperatures in combination with decreasing summer and increasing winter precipitation. The resulting overall drier conditions during the growing season will considerably impact forest growth there. We evaluate the consequences of increasing drought on the growth of the two locally most important broadleaf tree species common beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.). Three mixed forests of beech and oak were sampled along a west-east gradient of declining precipitation. In total we used 257 ring-width samples from 133 trees to build six species and site specific chronologies. Additionally, we modelled the soil water budget for each site. We performed continuous and discontinuous (pointer year) analysis of climate-tree-growth relationships with particular emphasis on inter-annual-variations and their dependence upon climatic factors (temperature, precipitation, soil moisture) and on the stability of the obtained relationships. Results of climate-growth correlations together with pointer year analysis indicate a strong dependency of growth of both species from water availability, especially during early summer (June and July). General correlation pattern between growth and climate are similar for both species, but climate sensitivity of beech is generally higher. We identified drought as the main driver of negative growth depressions in both species. Increasing drought stress along the gradient is expressed in higher correlations to climatic variables, higher sensitivity (variance) of growth, and a higher number of negative pointer years for both species. For beech we also found a significant trend of decreasing average growth rates along the gradient. Growth superiority of beech compared to oak declines with decreasing precipitation. The relationships were generally stable throughout the 20th century. A rise of sensitivity together with a higher frequency of negative pointer years during the last decades suggests that increasing climatic variability together with rising temperatures might be influencing growth of Fagus at the more humid sites. If we substitute space by time it seems that already small changes in precipitation regime can have considerable impact, especially on the growth of beech. Other, more drought tolerant species like oak might gain competitive advantages under the projected climatic changes.     

Adaptation of Asia-Pacific forests to climate change

Climate change is a threat to the stability and productivity of forest ecosystems throughout the Asia-Pacific region. The loss of forests due to climate-induced stress will have extensive adverse impacts on biodiversity and an array of ecosystem services that are essential for the maintenance of local economies and public health. Despite their importance, there is a lack of decision-support tools required to evaluate the potential effects of climate change on Asia-Pacific ecosystems and economies and to aid in the development of regionally appropriate adaptation and mitigation strategies. The project Adaptation of Asia-Pacific Forests to Climate Change, summarized herein, aims to address this lack of knowledge and tools and to provide support for regional managers to develop effective policy to increase the adaptive capacity of Asia-Pacific forest ecosystems. This objective has been achieved through the following activities: (1) development of a high-resolution climate downscaling model, ClimateAP, applicable to any location in the region; (2) development of climate niche models to evaluate how climate change might affect the distribution of suitable climatic conditions for regionally important tree species; (3) development and application of forest models to assess alternative management strategies in the context of management objectives and the projected impacts of climate change; (4) evaluation of models to assess forest fire risk and the relationship between forest fire and climate change; (5) development of a technique to assess ecosystem carbon storage using LiDAR; and (6) evaluation of how vegetation dynamics respond to climate change using remote sensing technology. All project outputs were developed with a focus on communication and extension to facilitate the dissemination of results to regional forest resource managers to support the development of effective mitigation and adaptation policy.     

Genetic variation of beech (Fagus sylvatica L.) in Rodopi (N.E. Greece)

The recent taxonomic classification of beech in Europe considers existence of one species (Fagus sylvatica L.) with two subspecies: F. sylvatica ssp. sylvatica and F. sylvatica ssp. orientalis. Four beech populations growing on the Greek part of the Rodopi Mountains were studied using morphological traits as well as DNA molecular markers (AFLPs and chloroplast DNA SSR). The aim of the study was to describe the variation patterns of beech in the Rodopi Mountains and to test the hypothesis of possible introgression between the beech subspecies’ sylvatica and orientalis in this area. Both morphological traits and gene markers revealed a possible influence of F. orientalis on the east side of Rodopi and at the low elevations, while characters resembling F. sylvatica were observed mainly on the western part of the mountains and in higher altitudes. There was a clinal increase of genetic diversity from the west to the east, reaching a level firstly reported for beech populations. These results can be explained either by the existence of a main refugial area for beech during the last glaciation or by the occurrence of a recent hybridization among the subspecies, which were spatially isolated during the last glaciation and came into reproductive contact during their postglacial remigration. These two scenarios are not necessarily mutually exclusive.