Dispersal limitation drives successional pathways in Central Siberian forests under current and intensified fire regimes

Fire is a primary driver of boreal forest dynamics. Intensifying fire regimes due to climate change may cause a shift in boreal forest composition toward reduced dominance of conifers and greater abundance of deciduous hardwoods, with potential biogeochemical and biophysical feedbacks to regional and global climate. This shift has already been observed in some North American boreal forests and has been attributed to changes in site conditions. However, it is unknown if the mechanisms controlling fire‐induced changes in deciduous hardwood cover are similar among different boreal forests, which differ in the ecological traits of the dominant tree species. To better understand the consequences of intensifying fire regimes in boreal forests, we studied postfire regeneration in five burns in the Central Siberian dark taiga, a vast but poorly studied boreal region. We combined field measurements, dendrochronological analysis, and seed‐source maps derived from high‐resolution satellite images to quantify the importance of site conditions (e.g., organic layer depth) vs. seed availability in shaping postfire regeneration. We show that dispersal limitation of evergreen conifers was the main factor determining postfire regeneration composition and density. Site conditions had significant but weaker effects. We used information on postfire regeneration to develop a classification scheme for successional pathways, representing the dominance of deciduous hardwoods vs. evergreen conifers at different successional stages. We estimated the spatial distribution of different successional pathways under alternative fire regime scenarios. Under intensified fire regimes, dispersal limitation of evergreen conifers is predicted to become more severe, primarily due to reduced abundance of surviving seed sources within burned areas. Increased dispersal limitation of evergreen conifers, in turn, is predicted to increase the prevalence of successional pathways dominated by deciduous hardwoods. The likely fire‐induced shift toward greater deciduous hardwood cover may affect climate–vegetation feedbacks via surface albedo, Bowen ratio, and carbon cycling.     

Leaf Mass per Area (LMA) and Its Relationship with Leaf Structure and Anatomy in 34 Mediterranean Woody Species along a Water Availability Gradient

Leaf mass per area (LMA) is a morphological trait widely used as a good indicator of plant functioning (i.e. photosynthetic and respiratory rates, chemical composition, resistance to herbivory, etc.). The LMA can be broken down into the leaf density (LD) and leaf volume to area ratio (LVA or thickness), which in turn are determined by anatomical tissues and chemical composition. The aim of this study is to understand the anatomical and chemical characteristics related to LMA variation in species growing in the field along a water availability gradient. We determined LMA and its components (LD, LVA and anatomical tissues) for 34 Mediterranean (20 evergreen and 14 deciduous) woody species. Variation in LMA was due to variation in both LD and LVA. For both deciduous and evergreen species LVA variation was strongly and positively related with mesophyll volume per area (VA or thickness), but for evergreen species positive relationships of LVA with the VA of epidermis, vascular plus sclerenchyma tissues and air spaces were found as well. The leaf carbon concentration was positively related with mesophyll VA in deciduous species, and with VA of vascular plus sclerenchymatic tissues in evergreens. Species occurring at the sites with lower water availability were generally characterised by a high LMA and LD.     

Ecophysiological and bioclimatic foundations for a global plant functional classification

Question: What plant properties might define plant functional types (PFTs) for the analysis of global vegetation responses to climate change, and what aspects of the physical environment might be expected to predict the distributions of PFTs? Methods: We review principles to explain the distribution of key plant traits as a function of bioclimatic variables. We focus on those whole‐plant and leaf traits that are commonly used to define biomes and PFTs in global maps and models. Results: Raunkiær's plant life forms (underlying most later classifications) describe different adaptive strategies for surviving low temperature or drought, while satisfying requirements for reproduction and growth. Simple conceptual models and published observations are used to quantify the adaptive significance of leaf size for temperature regulation, leaf consistency for maintaining transpiration under drought, and phenology for the optimization of annual carbon balance. A new compilation of experimental data supports the functional definition of tropical, warm‐temperate, temperate and boreal phanerophytes based on mechanisms for withstanding low temperature extremes. Chilling requirements are less well quantified, but are a necessary adjunct to cold tolerance. Functional traits generally confer both advantages and restrictions; the existence of trade‐offs contributes to the diversity of plants along bioclimatic gradients. Conclusions: Quantitative analysis of plant trait distributions against bioclimatic variables is becoming possible; this opens up new opportunities for PFT classification. A PFT classification based on bioclimatic responses will need to be enhanced by information on traits related to competition, successional dynamics and disturbance.     

Leaf dry matter content and lateral spread predict response to land use change for six subalpine grassland species

Question: Land‐use change has a major impact on terrestrial plant communities by affecting fertility and disturbance. We test how particular combinations of plant functional traits can predict species responses to these factors and their abundance in the field by examining whether trade‐offs at the trait level (fundamental trade‐offs) are linked to trade‐offs at the response level (secondary trade‐offs). Location: Central French Alps. Methods: We conducted a pot experiment in which we characterized plant trait syndromes by measuring whole plant and leaf traits for six dominant species, originating from contrasting subalpine grassland types. We characterized their response to nutrient availability, shading and clipping. We quantified factors linked with different land usage in the field to test the relevance of our experimental treatments. Results: We showed that land management affected nutrient concentration in soil, light availability and disturbance intensity. We identified particular suites of traits linked to plant stature and leaf structure which were associated with species responses to these environmental factors. Leaf dry matter content separates fast and slow growing species. Height and lateral spread separated tolerant and intolerant species to shade and clipping. Discussion and Conclusion: Two fundamental trade‐offs based on stature traits and leaf traits were linked to two secondary trade‐offs based on response to fertilization shade and mowing. Based on these trade‐offs, we discuss four different species strategies which could explain and predict species distributions and traits syndrome at community scale under different land‐uses in subalpine grasslands.     

Distinct Leaf‐trait Syndromes of Evergreen and Deciduous Trees in a Seasonally Dry Tropical Forest

In seasonally dry tropical forests, tree species can be deciduous, remaining without leaves throughout the dry season, or evergreen, retaining their leaves throughout the dry season. Deciduous and evergreen trees specialize in habitats that differ in water availability (hillside and riparian forest, respectively) and in their exposure to herbivore attack (seasonal and continuous, respectively). We asked whether syndromes of leaf traits in deciduous and evergreen trees were consistent with hypothesized abiotic and biotic selective pressures in their respective habitat. We measured seven leaf traits in 19 deciduous and 11 evergreen tree species in a dry tropical forest in Western Mexico, and measured rates of herbivory on 23 of these species. We investigated the covariance of leaf traits in syndromes related to phenology and associated physiology, and to anti‐herbivory defense. We found evidence for syndromes that separated phenological strategies among four traits: toughness, water content, specific leaf area, and carbon:nitrogen (C:N) ratios. We found a trade‐off between two other traits: trichomes and latex. Overall, evergreen species exhibited lower rates of herbivory than deciduous species. Lower rates of herbivory were explained by a syndrome of higher toughness, lower water content, and higher C:N ratios, which are traits representative of evergreen trees. Phenology and trait syndromes did not exhibit significant phylogenetic signal, consistent with the hypothesis of evolutionary convergence among phenologies and associated leaf‐trait syndromes. Our results suggest that deciduous and evergreen trees could respond to differential water availability and herbivory in their respective habitats by converging on distinct leaf‐trait syndromes. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Leaf trait co‐variation,response and effect in a chronosequence

Question: Is there any generality in terms of leaf trait correlations and the multiple role of leaf traits (response to and/or effect on) during secondary succession? Location: A secondary successional sere was sampled at four different ages since abandonment from several years to nearly 150 years on the Loess Plateau of northwestern China. Method: Specific leaf area (SLA), leaf mass per area (LMA), leaf nitrogen (N_mass, N_area), leaf phosphorus (P_mass, P_area) and leaf dry matter content (LDMC) were measured for all species recorded in the successional sere. Above‐ground net primary productivity (ANPP) and specific rate of litter mass loss (SRLML) were measured as surrogates for ecosystem properties. Soil total carbon (C) and nitrogen (N) were measured in each stage. Leaf traits were related to ecosystem properties and soil nutrient gradients, respectively. Results: LMA is correlated with N_area and P_area' and negatively with N_mass. Correlation between N_area and P_area was higher than between N_mass and P_mass. At the community level, field age, community hierarchy and their interaction explain 64.4 ‐ 93.5% of the variation in leaf traits. At the species level, field age explains 22.4 ‐ 45.5% of the variation in leaf traits (excl. P_area) while plant functional group has a significant effect only for N_mass. LDMC is correlated with ANPP and negatively with SRLML; P_mass is correlated with SRLML. Conclusions: Mean values of LMA, N_mass and N_area are close to the worldwide means, suggesting that large‐scale climate has a profound effect on leaf mass and leaf nitrogen allocation, while environmental gradients represented by succession have little influence on leaf‐trait values. Correlations between leaf traits, such as LMA‐N_area, LMA‐P_area and LMA‐N_mass shown in previous studies, are confirmed here. Although none of the leaf traits is proved to be both a response trait and an effect trait independent of time scale and community hierarchy, mass‐based leaf N is likely a sensitive response trait to soil C and N gradients. In addition, LDMC can be a marker for ANPP and SRLML, while mass‐based leaf P can be a marker for SRLML.     

缙云山森林土壤速效N、P、K时空特征研究

研究了缙云山森林生态系统内4个演替阶段群落的土壤速效N、P、K的时空特性,结果表明：①除灌草丛的速效K外,速效N、P、K含量在不同群落的土壤剖面上均具有明显的层次性,即腐殖质层（A）＞沉积层（B）＞母质层（C）。②A、B两层的速效N、K含量随群落演替方向升高,即灌草丛＜针叶林＜针阔混交林＜常绿阔叶林,速效P含量则为：针叶林＜针阔混交林＜灌草林＜常绿阔叶林。C层速效N、P、K含量似乎与群落演替规律无必然联系。③各群落内A、B层土壤速效N、P、K含量均表现出明显的季节动态,动态规律因元素类型和土壤层次而有差别。     

Periodicity of leaf growth and leaf dry mass changes in the evergreen and deciduous species of Southern Assam,India

The study described patterns of leaf dry mass change, leaf mass per area (LMA), relative growth rate and leaf life span (LL) for 14 evergreen and 7 deciduous species of a tropical forest of Southern Assam, India. Leaf expansion in both the groups was, in general, completed before June (i.e. well before the onset of monsoon rains). Although leaf dry mass during leaf initiation phase was significantly higher (P < 0.01) in evergreen species than in deciduous species, at the time of full leaf expansion, average leaf dry mass relative to the peak leaf dry mass, realised by the evergreen species was lower (66 %) than for deciduous species (76 %). Leaf dry mass increase in both groups continued after leaf full expansion. Evergreen species had a longer leaf dry mass steady phase than deciduous species (2–6 vs 2–3 months). Average LMA of mature leaves for evergreen species (77.43 g m^?2) was significantly greater than that of deciduous species (48.43 g m^?2). LL ranged from 165 days in Gmelina arborea (deciduous) to 509 days in Dipterocarpus turbinatus (evergreen). LMA was correlated positively with LL, indicating that evergreen species with higher leaf construction cost retain leaves for longer period to pay back. The average leaf dry mass loss before leaf shedding was greater (P < 0.01) for deciduous species (30.29 %) than for evergreen species (18.31 %). Although the cost of leaf construction in deciduous species was lower than for evergreen species, they replace leaves at a faster rate. Deciduous species perhaps compensate the cost involved in faster leaf replacement through higher reabsorption of dry mass during senescence, which they remobilise to initiate growth in the following spring when soil resources remain limiting.     

Responses of leaf structure and photosynthetic properties to intra-canopy light gradients: a common garden test with four broadleaf deciduous angiosperm and seven evergreen conifer tree species

Spectra of leaf traits in northern temperate forest canopies reflect major differences in leaf longevity between evergreen conifers and deciduous broadleaf angiosperms, as well as plastic modifications caused by within-crown shading. We investigated (1) whether long-lived conifer leaves exhibit similar intra-canopy plasticity as short-lived broadleaves, and (2) whether global interspecific relationships between photosynthesis, nitrogen, and leaf structure identified for sun leaves adequately describe leaves differentiated in response to light gradients. We studied structural and photosynthetic properties of intra-tree sun and shade foliage in adult trees of seven conifer and four broadleaf angiosperm species in a common garden in Poland. Shade leaves exhibited lower leaf mass-per-area (LMA) than sun leaves; however, the relative difference was smaller in conifers than in broadleaves. In broadleaves, LMA was correlated with lamina thickness and tissue density, while in conifers, it was correlated with thickness but not density. In broadleaves, but not in conifers, reduction of lamina thickness was correlated with a thinner palisade layer. The more conservative adjustment of conifer leaves could result from a combination of phylogenetic constraints, contrasting leaf anatomies and shoot geometries, but also from functional requirements of long-lived foliage. Mass-based nitrogen concentration (N(mass)) was similar between sun and shade leaves, and was lower in conifers than in deciduous broadleaved species. Given this, the smaller LMA in shade corresponded with a lower area-based N concentration (N(area)). In evergreen conifers, LMA and N(area) were less powerful predictors of area-based photosynthetic rate (A (max(area))) in comparison with deciduous broadleaved angiosperms. Multiple regression for sun and shade leaves showed that, in each group, A (max(mass)) was related to N(mass) but not to LMA, whereas LMA became a significant codeterminant of A (max(mass)) in analysis combining both groups. Thus, a fundamental mass-based relationship between photosynthesis, nitrogen, and leaf structure reported previously also exists in a dataset combining within-crown and across-functional type variation.     

A biome classification of China based on plant functional types and the BIOME3 model

A biome classification for China was established based on plant functional types (PFTs) using the BIOME3 model to include 16 biomes. In the eastern part of China, the PFTs of trees determine mostly the physiognomy of landscape. Biomes range from boreal deciduous coniferous forest/woodland, boreal mixed forest/woodland, temperate mixed forest, temperate broad-leaved deciduous forest, warm-temperate broad-leaved evergreen/mixed forest, warm-temperate/cool-temperate evergreen coniferous forest, xeric woodland/scrub, to tropical seasonal and rain forest, and tropical deciduous forest from north to south. In the northern and western part of China, grass is the dominant PFT. From northeast to west and southwest the biomes range from moist savannas, tall grassland, short grassland, dry savannas, arid shrubland/steppe, desert, to alpine tundra/ice/polar desert. Comparisons between the classification introduced here and the four classifications which were established over the past two decades, i.e. the vegetation classification, the vegetation division, the physical ecoregion, and the initial biome classification have showed that the different aims of biome classifications have resulted in different biome schemes each with its own unique characteristics and disadvantages for global change study. The new biome classification relies not only on climatic variables, but also on soil factor, vegetation functional variables, ecophysiological parameters and competition among the PFTs. It is a comprehensive classification that using multivariables better expresses the vegetation distribution and can be compared with world biome classifications. It can be easily used in the response study of Chinese biomes to global change, regionally and globally.     

Regional‐scale directional changes in abundance of tree species along a temperature gradient in Japan

Climate changes are assumed to shift the ranges of tree species and forest biomes. Such range shifts result from changes in abundances of tree species or functional types. Owing to global warming, the abundance of a tree species or functional type is expected to increase near the colder edge of its range and decrease near the warmer edge. This study examined directional changes in abundance and demographic parameters of forest trees along a temperature gradient, as well as a successional gradient, in Japan. Changes in the relative abundance of each of four functional types (evergreen broad‐leaved, deciduous broad‐leaved, evergreen temperate conifer, and evergreen boreal conifer) and the demography of each species (recruitment rate, mortality, and population growth rate) were analyzed in 39 permanent forest plots across the Japanese archipelago. Directional changes in the relative abundance of functional types were detected along the temperature gradient. Relative abundance of evergreen broad‐leaved trees increased near their colder range boundaries, especially in secondary forests, coinciding with the decrease in deciduous broad‐leaved trees. Similarly, relative abundance of deciduous broad‐leaved trees increased near their colder range boundaries, coinciding with the decrease in boreal conifers. These functional‐type‐level changes were mainly due to higher recruitment rates and partly to the lower mortality of individual species at colder sites. This is the first report to show that tree species abundances in temperate forests are changing directionally along a temperature gradient, which might be due to current or past climate changes as well as recovery from past disturbances.     

Climatic classification and ordination of the Spanish Sistema Central: relationships with potential vegetation

Comparisons among European, Japanese and North-American temperate deciduous woody floras revealed that there is no difference in shade-tolerance or in successional position between the compound- and simple-leaved species. Given that the compound-leaved species usually have greater biomass investments in non-productive throwaway supporting structures, it remained unclear how they could be as shade-tolerant as the simple-leaved analogues. To find out the role of the variability in leaf structure and composition in shade-tolerance of these species, foliar morphology and chemistry were analysed in 15 Estonian temperate compound-leaved deciduous woody taxa.Both foliar morphological and chemical parameters influenced the fractional investment of foliar biomass in petioles. The proportion of leaf biomass in petioles was independent of leaf size, but it increased with increasing leaflet number per leaf, suggesting that spacing rather than support requirements determined the biomass investment in petioles. The leaves with greater nitrogen concentrations also had larger foliar biomass investments in petioles. The latter effect possibly resulted from a greater water demand of functionally more active protein-rich leaves. Though the proportion of leaf biomass invested in petioles was high (for the whole material on average 15.9±0.4%), petioles were considerably cheaper to construct in terms of mineral nutrients than leaflets. e.g., petioles contained on average only 5.55±0.14% of total leaf nitrogen. Since in many cases the availability of mineral nutrients such as nitrogen rather than organic carbon sets limits to total leaf biomass on the plant, I suggested, contrary to previous claims, that the costs for foliage formation should not necessarily be different between compound- and simple-leaved species. Compound-leaved species also fit the basic relationships previously observed in simple-leaved analogues. Leaf size increased and leaf dry mass per area (LMA) decreased with increasing shade-tolerance. Thus, more shade-tolerant species construct a more effective foliar display for light interception at low irradiance with similar biomass investment in leaves. Species shade-tolerance was independent of biomass investment in petioles. However, due to the genotypic plasticity in LMA, more shade-tolerant species supported more foliar area at a constant leaf biomass investment in petioles.     

Evolution of the leaf economics spectrum in herbs: Evidence from environmental divergences in leaf physiology across Helianthus (Asteraceae)

The leaf economics spectrum (LES) describes a major axis of plant functional trait variation worldwide, defining suites of leaf traits aligned with resource‐acquisitive to resource‐conservative ecological strategies. The LES has been interpreted to arise from leaf‐level trade‐offs among ecophysiological traits common to all plants. However, it has been suggested that the defining leaf‐level trade‐offs of the LES may not hold within specific functional groups (e.g., herbs) nor within many groups of closely related species, which challenges the usefulness of the LES paradigm across evolutionary scales. Here, we examine the evolution of the LES across 28 species of the diverse herbaceous genus Helianthus (the sunflowers), which occupies a wide range of habitats and climate variation across North America. Using a phylogenetic comparative approach, we find repeated evolution of more resource‐acquisitive LES strategies in cooler, drier, and more fertile environments. We also find macroevolutionary correlations among LES traits that recapitulate aspects of the global LES, but with one major difference: leaf mass per area is uncorrelated with leaf lifespan. This indicates that whole‐plant processes likely drive variation in leaf lifespan across Helianthus, rather than leaf‐level trade‐offs. These results suggest that LES patterns do not reflect universal physiological trade‐offs at small evolutionary scales.     

Leaf functional traits vary with the adult height of plant species in forest communities

Aims Within-community variation accounts for a remarkable proportion of the variation in leaf functional traits. Plant height may be used to explain within-community variances of leaf traits because different microenvironments, especially light intensity, may occur at different heights. This study determines whether or not leaf nitrogen (N) and phosphorus (P) contents as well as leaf mass per area (LMA) are interspecifically correlated with the adult height of forest woody species. We also discuss these relationships with respect to community structure and functions of the ecosystem.Methods A total of 136 dicotyledonous woody species from 6 natural forests (3 evergreen and 3 deciduous ones) in East China (18°44′–45°25′N, 108°50′–128°05′E) were investigated. For each of the 157 species–site combinations, 6 traits were measured: plant adult height relative to the forest canopy (H R), leaf N and P contents per unit area (N area and P area), N and P contents per unit dry mass (N mass and P mass) and LMA. The total variances of each leaf trait across sites were partitioned in a hierarchical manner. The relationships between leaf traits and H R within forest communities were then analyzed using both standardized major axis regression and Felsenstein's phylogenetic independent contrasts. Relationships between evergreen and deciduous forests were compared by linear mixed models.Important findings H R is a robust predictor of leaf N area, P area and LMA, explaining 36.7, 39.4 and 12.0% of their total variations across forests, respectively. Leaf N area, P area and LMA increased with H R in all of the studied forests, with slopes that were steeper in evergreen forests than in deciduous ones. Leaf N mass and P mass showed no significant relationship with H R generally. The increase in leaf N area, P area and LMA with H R across species is assumed to maximize community photosynthesis and may favor species with larger H R .     

Photosynthesis or persistence: nitrogen allocation in leaves of evergreen and deciduous Quercus species

Photosynthetic nitrogen use efficiency (PNUE, photosynthetic capacity per unit leaf nitrogen) is one of the most important factors for the interspecific variation in photosynthetic capacity. PNUE was analysed in two evergreen and two deciduous species of the genus Quercus. PNUE was lower in evergreen than in deciduous species, which was primarily ascribed to a smaller fraction of nitrogen allocated to the photosynthetic apparatus in evergreen species. Leaf nitrogen was further analysed into proteins in the water‐soluble, the detergent‐soluble, and the detergent‐insoluble fractions. It was assumed that the detergent‐insoluble protein represented the cell wall proteins. The fraction of nitrogen allocated to the detergent‐insoluble protein was greater in evergreen than in deciduous leaves. Thus the smaller allocation of nitrogen to the photosynthetic apparatus in evergreen species was associated with the greater allocation to cell walls. Across species, the fraction of nitrogen in detergent‐insoluble proteins was positively correlated with leaf mass per area, whereas that in the photosynthetic proteins was negatively correlated. There may be a trade‐off in nitrogen partitioning between components pertaining to productivity (photosynthetic proteins) and those pertaining to persistence (structural proteins). This trade‐off may result in the convergence of leaf traits, where species with a longer leaf life‐span have a greater leaf mass per area, lower photosynthetic capacity, and lower PNUE regardless of life form, phyllogeny, and biome.     

Environmental heterogeneity and spatiotemporal variability in plant defense traits

At the scale of the local plant community, we know very little about how spatial and temporal environmental heterogeneity affects the diversity in types and levels of plant defenses. If environmental heterogeneity is an important mechanism influencing plant defense traits, then defense expression should co‐vary spatially with environmental conditions and change as succession progresses. In this study, we examined how spatial heterogeneity and succession influence putative resistance and tolerance trait levels in late goldenrod Solidago altissima. We quantified the spatial distributions of herbivore damage and traits associated with resistance (leaf toughness, phenolics), tolerance (specific leaf area, relative growth rate, leaf addition rate and leaf senescence rate), and fitness (height, diameter, inflorescence biomass) of goldenrods within replicate early‐ and late‐successional fields. Also, we characterized the local neighborhood (stem density, canopy cover, ground vegetative cover) and edaphic conditions (soil moisture, pH, N) surrounding each target ramet, and determined relationships between these environmental variables and goldenrod trait levels. The distribution of traits within fields was strongly non‐random, and defense‐trait levels were more strongly spatially structured (i.e. autocorrelated) in late‐ than in early‐successional fields. Also, defense traits were most strongly correlated with aspects of the local plant neighborhood, and these relationships differed in important ways between successional stages. In late‐successional fields, tolerance trait specific leaf area was positively correlated with canopy cover and negatively correlated with stem density. In early‐successional fields, the relationship between ground vegetative cover and resistance (i.e. 1 – damage) was significantly stronger than in late‐successional fields. A novel insight from this study is the possibility that changes in the biotic environment during succession may shift the expression of defense from a resistance to a tolerance strategy in our system. This study highlights the context dependence of plant defense trait levels, which may promote their spatial and temporal variability in heterogeneous landscapes.     

Diversity of phenolic compounds and plant traits in coexisting Patagonian desert shrub species of Argentina

Desert shrubs often accumulate different types of phenolic compounds but what determines the amount and diversity of these compounds is an issue scarcely explored. The aim of this study was to assess differences in the amount and diversity of phenolic compounds in leaves among coexisting shrub species differing in rooting depth and leaf turnover. We hypothesized that the diversity and amount of phenolic compounds in leaves of desert shrubs are related to access to soil water through rooting depth, and to leaf turnover. The study was carried out in the Patagonian Monte of Argentina. We collected green leaves of six species representing the dominant shrub morphotypes (tall evergreen, tall deciduous, and medium evergreen shrubs) and assessed lignin concentration and groups of soluble phenols obtained by sequential extraction with ethyl ether, ethyl acetate, and amyl alcohol. We also assessed nitrogen concentration in leaves and leaf mass per unit area (LMA) as traits related to leaf lifespan. The diversity of phenolic compounds was higher in green leaves of tall shrubs with deep rooting depth than in those of medium evergreen shrubs with shallow rooting depth. Diversity of phenolic compounds in green leaves was negatively related to lignin concentration. Evergreen shrubs had higher amount of phenolic compounds in green leaves than deciduous ones and the total amount of phenolic compounds in green leaves was positively related to LMA. We concluded that access to soil water sources and leaf turnover were related to the amount and diversity of phenolic compounds in green leaves of desert shrub species and these results are consistent with those predicted by the resource availability theory for plants from resource-rich and resource-poor habitats.     

Spatial patterns of photosynthetic characteristics and leaf physical traits of plants in the Loess Plateau of China

The spatial patterns of photosynthetic characteristics and leaf physical traits of 171 plants belonging to nine life-forms or functional groups (trees, shrubs, herbs, evergreen trees, deciduous trees, C₃ and C₄ herbaceous plants, leguminous and non-leguminous species) and their relationships with environmental factors in seven sites, Yangling, Yongshou, Tongchuan, Fuxian, Ansai, Mizhi and Shenmu, ranging from south to north in the Loess Plateau of China were studied. The results showed that the leaf light-saturated photosynthetic rate (P_max), photosynthetic nitrogen use efficiency (PNUE), chlorophyll content (Chl), and leaf mass per area (LMA) of all the plants in the Loess Plateau varied significantly among three life-form groups, i.e., trees, shrubs and herbs, and two groups, i.e., evergreen trees and deciduous trees, but leaf nitrogen content differed little among different life-form groups. For the 171 plants in the Loess Plateau, leaf P_max was positively correlated with PNUE. The leaf nitrogen content per unit area (N_area) was positively correlated but Chl was negatively correlated with the LMA. When controlling the LMA, the N_area was positively correlated with the Chl (partial r = 0.20, P < 0.05). With regard to relationships between photosynthetic characteristics and leaf physical traits, the P_max was positively correlated with N _area, while the PNUE was positively correlated with the Chl and negatively correlated with the N_area and LMA. For all the species in the Loess Plateau, the PNUE was negatively correlated with the latitude and annual solar radiation (ASR), but positively correlated with the mean annual rainfall (MAR) and mean annual temperature (MAT). With regard to the leaf physical traits, the leaf Chl was negatively correlated with the latitude and ASR, but positively correlated with the MAR and MAT. However, the N_area and LMA were positively correlated with the latitude and ASR, but negatively correlated with the MAR and MAT. In general, leaf N_area and LMA increased, while PNUE and Chl decreased with increases in the latitude and ASR and decreases in MAR and MAT. Electronic supplementary material The online version of this article () contains supplementary material, which is available to authorized users.     

Micro-site performance of evergreen and deciduous dwarf shrubs in a subarctic heath in relation to nitrogen status

The relationship between micro-site nitrogen status (total soil N conc., total vegetation N, and leaf N conc.) and performance (biomass) for two evergreen and two deciduous dwarf shrubs was studied in a subarctic heath. One deciduous species, Vaccinium uliginosum , dominated on sites having the highest vegetation nitrogen content and highest leaf nitrogen concentrations. The evergreen Empetrum hermaphroditum dominated on sites with a low total vegetation nitrogen content and low leaf nitrogen concentrations. Two other species, V. myrtillus (deciduous) and V. vitis-idaea (evergreen), showed intermediate patterns that were not clearly separated from either V. uliginosum or Empetrum . Soil nitrogen content showed no significant differences between micro-sites. Possible physiological reasons for the observed patterns are discussed.     

后河自然保护区常绿落叶阔叶混交林群落学特征研究

研究了后河国家级自然保护区常绿落叶阔叶混交林的区系特点、群落学特征。结果表明 :后河国家级自然保护区常绿落叶阔叶混交林群落物种丰富 ,有 110科 2 88属 5 34种 ,科的分布型以热带类型为主 ,属的分布型温带类型占优势 ,反映该群落区系以温带为主的亚热带区系特点 ;该混交林主要由以单叶、中型叶及非全缘叶为主的常绿、落叶阔叶高位芽植物组成 ,群落层次分明 ,从上至下 ,落叶阔叶树种所占比例逐渐减少 ,常绿阔叶树种比例逐渐增加 ;除两低海拔样地外 ,多样性变化幅度较小 ,各样地的物种多样性指数变化趋势基本一致。