How tree species identity and diversity affect light transmittance to the understory in mature temperate forests

Light is a key resource for plant growth and is of particular importance in forest ecosystems, because of the strong vertical structure leading to successive light interception from canopy to forest floor. Tree species differ in the quantity and heterogeneity of light they transmit. We expect decreases in both the quantity and spatial heterogeneity of light transmittance in mixed stands relative to monocultures, due to complementarity effects and niche filling. We tested the degree to which tree species identity and diversity affected, via differences in tree and shrub cover, the spatiotemporal variation in light availability before, during, and after leaf expansion. Plots with different combinations of three tree species with contrasting light transmittance were selected to obtain a diversity gradient from monocultures to three species mixtures. Light transmittance to the forest floor was measured with hemispherical photography. Increased tree diversity led to increased canopy packing and decreased spatial light heterogeneity at the forest floor in all of the time periods. During leaf expansion, light transmittance did differ between the different tree species and timing of leaf expansion might thus be an important source of variation in light regimes for understory plant species. Although light transmittance at the canopy level after leaf expansion was not measured directly, it most likely differed between tree species and decreased in mixtures due to canopy packing. A complementary shrub layer led, however, to similar light levels at the forest floor in all species combinations in our plots. Synthesis. We find that a complementary shrub layer exploits the higher light availability in particular tree species combinations. Resources at the forest floor are thus ultimately determined by the combined effect of the tree and shrub layer. Mixing species led to less heterogeneity in the amount of light, reducing abiotic niche variability.     

Soil phosphorus heterogeneity promotes tree species diversity and phylogenetic clustering in a tropical seasonal rainforest

The niche theory predicts that environmental heterogeneity and species diversity are positively correlated in tropical forests, whereas the neutral theory suggests that stochastic processes are more important in determining species diversity. This study sought to investigate the effects of soil nutrient (nitrogen and phosphorus) heterogeneity on tree species diversity in the Xishuangbanna tropical seasonal rainforest in southwestern China. Thirty‐nine plots of 400 m² (20 × 20 m) were randomly located in the Xishuangbanna tropical seasonal rainforest. Within each plot, soil nutrient (nitrogen and phosphorus) availability and heterogeneity, tree species diversity, and community phylogenetic structure were measured. Soil phosphorus heterogeneity and tree species diversity in each plot were positively correlated, while phosphorus availability and tree species diversity were not. The trees in plots with low soil phosphorus heterogeneity were phylogenetically overdispersed, while the phylogenetic structure of trees within the plots became clustered as heterogeneity increased. Neither nitrogen availability nor its heterogeneity was correlated to tree species diversity or the phylogenetic structure of trees within the plots. The interspecific competition in the forest plots with low soil phosphorus heterogeneity could lead to an overdispersed community. However, as heterogeneity increase, more closely related species may be able to coexist together and lead to a clustered community. Our results indicate that soil phosphorus heterogeneity significantly affects tree diversity in the Xishuangbanna tropical seasonal rainforest, suggesting that deterministic processes are dominant in this tropical forest assembly.     

Correlations of understory herb distribution patterns with microhabitats under different tree species in a mixed mesophytic forest

Summary This study examines the role of canopy trees in the formation and maintenance of different herb microhabitats in a mixed mesophytic forest stand. Herb abundance and reproductive success were recorded in 54 circular plots under seven species of canopy trees and in 15 circular control plots>2 m from any tree. Soil moisture, soil nutrient levels, litter depth, and light intensity were measured in a subset of these plots. Ordination of plots by both herb relative abundance and by reproductive success of common species indicated that herb assemblages under most canopy tree species were similar to those away from trees. However, herb assemblages under Fagus grandifolia trees differed moderately from the others while plots under Quercus alba trees supported significantly different herb assemblages. Analyses of variance revealed that several herb species occurred at significantly closer mean distance to the base of Q. alba or Fagus trees or at higher densities under these tree species. Soils around Q. alba trees had significantly higher concentrations of calcium and sulfate ions, and higher pH than plots under other tree species and control plots. This correlated closely with Q. alba stemflow which had higher concentrations of calcium and sulfate ions and lower concentrations of hydrogen ions than stemflow from other trees at this site. The slightly lower soil pH near the base of Fagus trees may have been related to the high volumes of stemflow produced by this species. Stepwise regression showed significant correlations between abundances of five common herb species and soil nutrient patterns. Maintenance of spatial heterogeneity in forest floor resources by the presence of different species of canopy trees may therefore be important in the maintenance of diversity in these understory herb communities.     

The influence of tree species on canopy soil nutrient status in a tropical lowland wet forest in Costa Rica

The canopy is host to a large percentage of the flora and fauna in tropical wet forests and is distinct from the forest floor in plant richness, soil type and microclimate. In this study, we examined the influence of tree species and season on soil nutrient cycling processes in canopy soils of four tree species common to Costa Rican wet forests. We also compared the canopy soils to the associated forest floor mineral soils. Both tree species and season had strong effects on canopy soil nutrients and processes. Canopy soils from trees with high litter lignin concentrations had higher net N-mineralization rates and higher dissolved inorganic N concentrations than those with low lignin concentrations. During the dry season, net N-immobilization occurred and dissolved organic and inorganic N and available P concentrations were significantly higher than during the wet season. Overall, canopy soils had higher N levels and higher fungi + bacteria richness than forest floor mineral soils. The differences in canopy soil properties observed among tree species indicates that these species have distinct N cycles that reflect differences in both soil origin and biological controls.     

The effects of stand structure on ground-floor bryophyte assemblages in temperate mixed forests

The effect of tree species composition, stand structure characteristics and substrate availability on ground-floor bryophyte assemblages was studied in mixed deciduous forests of Western Hungary. Species composition, species richness and cover of bryophytes occurring on the soil and logs were analysed as dependent variables. The whole assemblage and functional groups defined on the basis of substrate preference were investigated separately. Substrate availability (open soil, logs) was the most prominent factor in determining species composition, cover and diversity positively, while the litter of deciduous trees had a negative effect on the occurrence of forest floor bryophytes. Besides, bryophyte species richness increased with tree species and stand structural diversity, and for specialist epiphytic and epixylic species log volume was essential. Sapling density and light heterogeneity were influential on bryophyte cover, especially for the dominant terricolous species. Many variables of the forest floor bryophyte community can be estimated efficiently by examining stand structure in the studied region. Selective cutting increasing tree species diversity, stand structural heterogeneity and dead wood volume can maintain higher bryophyte diversity in this region than the shelter-wood system producing even-aged, monodominant, structurally homogenous stands.     

Spatial heterogeneity of understory vegetation and soil in an Alaskan upland boreal forest fire chronosequence

In this study we characterized spatial heterogeneity of soil carbon and nitrogen pools, soil moisture, and soil pH of the first 15?cm of the soil profile; depth of the organic horizon; forest floor covers; and understory vegetation abundances in three sites (1999, 1987 and 1920 wildfires) of a boreal forest chronosequence of interior Alaska. We also investigated the cross-dependence between understory vegetation distribution and soil characteristics. Our results showed higher microbial respiration rates and microbial biomass in the oldest site and greater net N mineralization rates in the mid-successional site. Although spatial heterogeneity was absent at the scale studied for the majority of soil variables (60%), understory vegetation abundances and forest floor cover, spatial heterogeneity decreased with time after fire for the depth of organic horizon, soil microbial biomass, N mineralization rates and feathermoss cover. Our results also showed that increasing time after fire decreased the number of correlations between understory vegetation and soil characteristics while it increased between forest floor covers and soil characteristics. Overall, our study suggest that fire initially creates a patchy mosaic of forest floor cover, from fire hot spots, where high intensity burning exposes mineral soil, to practically unburned areas with intact mosses and lichens. As time since fire passes, forest floor cover and soil characteristics tend to become more uniform as understory species fill in severely burned areas.     

Spatiotemporal changes of beetle communities across a tree diversity gradient

Aim Plant and arthropod diversity are often related, but data on the role of mature tree diversity on canopy insect communities are fragmentary. We compare species richness of canopy beetles across a tree diversity gradient ranging from mono‐dominant beech to mixed stands within a deciduous forest, and analyse community composition changes across space and time. Location Germany’s largest exclusively deciduous forest, the Hainich National Park (Thuringia). Methods We used flight interception traps to assess the beetle fauna of various tree species, and applied additive partitioning to examine spatiotemporal patterns of diversity. Results Species richness of beetle communities increased across the tree diversity gradient from 99 to 181 species per forest stand. Intra‐ and interspecific spatial turnover among trees contributed more than temporal turnover among months to the total γ‐beetle diversity of the sampled stands. However, due to parallel increases in the number of habitat generalists and the number of species in each feeding guild (herbivores, predators and fungivores), no proportional changes in community composition could be observed. If only beech trees were analysed across the gradient, patterns were similar but temporal (monthly) species turnover was higher compared to spatial turnover among trees and not related to tree diversity. Main conclusions The changes in species richness and community composition across the gradient can be explained by habitat heterogeneity, which increased with the mix of tree species. We conclude that understanding temporal and spatial species turnover is the key to understanding biodiversity patterns. Mono‐dominant beech stands are insufficient to conserve fully the regional species richness of the remaining semi‐natural deciduous forest habitats in Central Europe, and analysing beech alone would have resulted in the misleading conclusion that temporal (monthly) turnover contributes more to beetle diversity than spatial turnover among different tree species or tree individuals.     

Intermediary disturbance increases tree diversity in riverine forest of southern Brazil

Floods are frequently associated with disturbance in structuring riverine forests and they lead to environmental heterogeneity over space and time. We evaluated the distribution of tree species, ecological groups, species richness and diversity from the point bar to the slope of a riverside forest in southern Brazil (Lat. 30°01′S, Long. 52°47′W) to analyze the effects of flooding on soil properties and forest structure. A plot of 50 × 200 m divided in five contiguous transects of 10 × 200 m parallel to the river was installed, where we measured all the individual trees with pbh ≥ 15 cm. A detailed topographical and soil survey was carried out across the plot and indicated significant differences in organic matter and most mineral nutrients through the topographical gradient. The 1,229 surveyed individuals belonged to 72 species and 35 families. We used Partial CCA and Species Indicator Analysis to observe the spatial distribution of species. Both analyses showed that species distribution was strongly related to the flooding gradient, soil properties and also by space and pure spatial structuring of species and environmental variables (spatial autocorrelation), although a large part of variation remains unexplained. The ecological groups of forest stratification, plant dispersal and requirements for germination indicated slight differences among frequently, occasional and non-flooded transects. Species richness and diversity were higher at intermediate elevations and were associated to the increased spatial–temporal environmental heterogeneity. Across the plot, the direct influence of flooding on tree species distribution created a vegetation zonation that is determined by predicted ecological traits.     

Functional Composition of Tree Communities Changed Topsoil Properties in an Old Experimental Tropical Plantation

Forest biogeochemistry is strongly determined by the interaction between the tree community and the topsoil. Functional strategies of tree species are coupled to specific chemical leaf traits, and thus also to litter composition, which affects mineral soil characteristics. The limited understanding on this interaction is mainly based on shorter-term common garden experiments in temperate forest, and needs to be extended to other forest types and climates if we want to understand the universality of this linkage. In particular, for highly diverse tropical forests, our understanding of this interaction remains limited. Using an old experimental plantation within the central Congo basin, we examined the relationship between leaf and litter chemical composition and topsoil properties. Canopy, litter and topsoil characteristics were measured and we determined how the community-level leaf and litter chemical composition altered the topsoil carbon, major plant nutrients and exchangeable cation concentration, acidity and pH over the last eight decades. We found that functional composition strongly affected topsoil pH. In turn, topsoil pH strongly determined the soil total carbon and available phosphorus, total nitrogen and exchangeable potassium. Our results indicate that, as observed in temperate common garden experiments, trees alter chemical topsoil properties primarily through soil acidification, differently induced by functional composition of the tree community. The strong link between this community-level composition and topsoil characteristics, on a highly representative soil type for the tropics, improves our understanding of tropical forests biogeochemistry.     

LiDAR GEDI derived tree canopy height heterogeneity reveals patterns of biodiversity in forest ecosystems

The “Height Variation Hypothesis” is an indirect approach used to estimate forest biodiversity through remote sensing data, stating that greater tree height heterogeneity (HH) measured by CHM LiDAR data indicates higher forest structure complexity and tree species diversity. This approach has traditionally been analyzed using only airborne LiDAR data, which limits its application to the availability of the dedicated flight campaigns. In this study we analyzed the relationship between tree species diversity and HH, calculated with four different heterogeneity indices using two freely available CHMs derived from the new space-borne GEDI LiDAR data. The first, with a spatial resolution of 30 m, was produced through a regression tree machine learning algorithm integrating GEDI LiDAR data and Landsat optical information. The second, with a spatial resolution of 10 m, was created using Sentinel-2 images and a deep learning convolutional neural network. We tested this approach separately in 30 forest plots situated in the northern Italian Alps, in 100 plots in the forested area of Traunstein (Germany) and successively in all the 130 plots through a cross-validation analysis. Forest density information was also included as influencing factor in a multiple regression analysis. Our results show that the GEDI CHMs can be used to assess biodiversity patterns in forest ecosystems through the estimation of the HH that is correlated to the tree species diversity. However, the results also indicate that this method is influenced by different factors including the GEDI CHMs dataset of choice and their related spatial resolution, the heterogeneity indices used to calculate the HH and the forest density. Our finding suggest that GEDI LIDAR data can be a valuable tool in the estimation of forest tree heterogeneity and related tree species diversity in forest ecosystems, which can aid in global biodiversity estimation.     

西双版纳热带季节雨林土壤养分空间异质性对乔木树种多样性的影响

生态位理论认为,养分空间异质性分布会减少种间竞争排斥而有助于物种共存;而中性理论则认为群落树种呈独立于环境特征的随机分布。为研究土壤养分异质性与乔木树种多样性的联系,在西双版纳热带季节雨林随机设置了16个1 hm~2样方,调查了各样方乔木树种多样性,计算了各样方土壤有效氮(AN)、有效磷(EP)、有效钾(EK)、有机质(OM)、p H、总氮(TN)、总磷(TP)、总钾(TK)的变异系数以代表各样方土壤养分空间异质性分布的相对水平。Pearson相关分析表明,TK变异系数与乔木树种丰富度、Shannon-wiener指数呈显著正相关(P0.05),表明该区域不同植物可能在钾资源的利用上存在明显的生态位分化,钾的异质性有助于树种共存;OM、AN变异系数与Pielou均匀度指数呈明显正相关(P0.1),在一定程度上说明了这些养分的空间异质性缓解了种间竞争压力,树种多度分布相对均匀,有助于树种共存。除TK外,其它土壤指标的变异系数与乔木树种多样性的正相关性均不显著(P0.05),表明这些养分的空间异质性分布对乔木树种多样性的影响相对较小,中性或其它生态学过程可能掩盖了这些养分的空间异质性分布对乔木树种多样性的影响。这说明,土壤养分空间异质性可能在一定程度上促进了树种共存,但同时应当重视中性过程等在西双版纳热带雨林群落构建中的作用。     

Leaf litter inputs reinforce islands of nitrogen fertility in a lowland tropical forest

The role of lowland tropical forest tree communities in shaping soil nutrient cycling has been challenging to elucidate in the face of high species diversity. Previously, we showed that differences in tree species composition and canopy foliar nitrogen (N) concentrations correlated with differences in soil N availability in a mature Costa Rican rainforest. Here, we investigate potential mechanisms explaining this correlation. We used imaging spectroscopy to identify study plots containing 10–20 canopy trees with either high or low mean canopy N relative to the landscape mean. Plots were restricted to an uplifted terrace with relatively uniform parent material and climate. In order to assess whether canopy and soil N could be linked by litterfall inputs, we tracked litter production in the plots and measured rates of litter decay and the carbon and N content of leaf litter and leaf litter leachate. We also compared the abundance of putative N fixing trees and rates of free-living N fixation as well as soil pH, texture, cation exchange capacity, and topographic curvature to assess whether biological N fixation and/or soil properties could account for differences in soil N that were, in turn, imprinted on the canopy. We found no evidence of differences in legume communities, free-living N fixation, or abiotic properties. However, soils beneath high canopy N assemblages received ~ 60% more N via leaf litterfall due to variability in litter N content between plot types. The correlation of N in canopy leaves, leaf litter, and soil suggests that, under similar abiotic conditions, litterfall-mediated feedbacks can help maintain soil N differences among tropical tree assemblages in this diverse tropical forest.

     

Fungal disease incidence along tree diversity gradients depends on latitude in European forests

European forests host a diversity of tree species that are increasingly threatened by fungal pathogens, which may have cascading consequences for forest ecosystems and their functioning. Previous experimental studies suggest that foliar and root pathogen abundance and disease severity decrease with increasing tree species diversity, but evidences from natural forests are rare. Here, we tested whether foliar fungal disease incidence was negatively affected by tree species diversity in different forest types across Europe. We measured the foliar fungal disease incidence on 16 different tree species in 209 plots in six European countries, representing a forest‐type gradient from the Mediterranean to boreal forests. Forest plots of single species (monoculture plots) and those with different combinations of two to five tree species (mixed species plots) were compared. Specifically, we analyzed the influence of tree species richness, functional type (conifer vs. broadleaved) and phylogenetic diversity on overall fungal disease incidence. The effect of tree species richness on disease incidence varied with latitude and functional type. Disease incidence tended to increase with tree diversity, in particular in northern latitudes. Disease incidence decreased with tree species richness in conifers, but not in broadleaved trees. However, for specific damage symptoms, no tree species richness effects were observed. Although the patterns were weak, susceptibility of forests to disease appears to depend on the forest site and tree type.     

Vertical gradients in leaf trait diversity in a New Zealand forest

Leaves come in a remarkable diversity of sizes and shapes. However, spatial patterns in leaf trait diversity are rarely investigated and poorly resolved. We used a hierarchical approach to evaluate vertical variability in leaf morphology (i.e., leaf trait diversity) in 16 common tree and shrub species inhabiting a New Zealand forest. Height-related heterogeneity in leaf area, specific leaf area, circularity and length to width ratio was analyzed at three scales: (1) among leaves within plants, (2) among plants within species and (3) among species within functional groups (i.e., trees vs. shrubs). Results were scale dependent. Among-leaf morphological diversity was unrelated to plant height. Among-individual morphological diversity increased with the average height of each species, indicating that taller plant species express a greater range of leaf traits than shorter species. Among-species morphological diversity was higher in shrubs than in trees. We hypothesize that scale-dependent patterns in leaf trait diversity result from scale-dependent adaptations to forest environmental conditions. As trees grow from the forest floor into the canopy, they are exposed to a range of environmental conditions, which may select for a range of leaf traits through ontogeny. Conversely, shrubs never reach the forest canopy and may instead be differentially adapted to suites of environmental conditions associated with different stages of forest recovery from tree-fall disturbances. Overall results indicate that vertical patterns in leaf trait diversity exist. However, their strength and directionality are strongly scale-dependent, suggesting that different processes govern leaf shape diversity at different levels of ecological organization.     

Restoration of Tropical Moist Forests on Bauxite-Mined Lands in the Brazilian Amazon

We evaluated forest structure and composition in 9- to 13-year-old stands established on a bauxite-mined site at Trombetas (Pará), Brazil, using four different reforestation techniques following initial site preparation and topsoil replacement. These techniques included reliance on natural forest regeneration, mixed commercial species plantings of mostly exotic timber trees, direct seeding with mostly native early successional tree species, and mixed native species plantings of more than 70 tree species (the current operational restoration treatment at this site). Replicated fixed-radius plots in each treatment and in undisturbed primary forest were used to quantify the canopy and understory structure and the abundance and diversity of all vascular plant species. Treatment comparisons considered regeneration density, species richness and diversity for all floristic categories, and, for trees and shrubs, the relative contribution of initial planting and subsequent regeneration from soil seed banks and seed inputs from nearby primary forests. With the possible exception of the stands of mixed commercial species, which were superior to all others in terms of tree basal-area development but relatively poor in species richness, all treatments were structurally and floristically diverse, with a high probability of long-term restoration success. Of these, the mixed native species plantings appeared to be at least risk of arrested succession due to the dominance of a broader range of tree species of different successional stages or expected life spans. In all treatments, several locally important families of primary forest trees (Annonaceae, Chrysobalanaceae, Lauraceae, Palmae and Sapotaceae) were markedly underrepresented due to a combination of poor survival of initial plantings and limitations on seed dispersal from the surrounding primary forest.     

Influence of tree hollow characteristics and forest structure on saproxylic beetle diversity in tree hollows in managed forests in a regional comparison

Tree hollows are among the rarest habitats in today''s Central European managed forests but are considered key structures for high biodiversity in forests. To analyze and compare the effects of tree hollow characteristics and forest structure on diversity of saproxylic beetles in tree hollows in differently structured managed forests, we examined between 41 and 50 tree hollows in beech trees in each of three state forest management districts in Germany. During the two‐year study, we collected 283 saproxylic beetle species (5880 individuals; 22% threatened species), using emergence traps. At small spatial scales, the size of hollow entrance and the number of surrounding microhabitat structures positively influenced beetle diversity, while the stage of wood mould decomposition had a negative influence, across all three forest districts. We utilized forest inventory data to analyze the effects of forest structure in radii of 50–500 m around tree hollows on saproxylic beetle diversity in the hollows. At these larger spatial scales, the three forest management districts differed remarkably regarding the parameters that influenced saproxylic beetle diversity in tree hollows. In Ebrach, characterized by mostly deciduous trees, the amount of dead wood positively influenced beetle diversity. In the mostly coniferous Fichtelberg forest district, with highly isolated tree hollows, in contrast, only the proportion of beech trees around the focal tree hollows showed a positive influence on beetle diversity. In Kelheim, characterized by mixed forest stands, there were no significant relationships between forest structure and beetle diversity in tree hollows. In this study, the same local tree hollow parameters influenced saproxylic beetle diversity in all three study regions, while parameters of forest structure at larger spatial scales differed in their importance, depending on tree‐species composition.     

Point patterns of tree distribution determined by habitat heterogeneity and dispersal limitation

Understanding processes underlying spatial distribution of tree species is fundamental to studying species coexistence and diversity. This study modeled point patterns of tree distribution, expressed by Cartesian coordinates of individual trees within a mapped forest stand, for the purpose of identifying processes that may generate spatial patterns of tree communities. We used four primary point pattern processes (homogeneous Poisson process, inhomogeneous Poisson process, homogeneous Thomas process, and inhomogeneous Thomas process) to model tree distribution in two stem-mapped forests in Taiwan, Republic of China. These four models simulate spatial processes of habitat association and seed dispersal, allowing us to evaluate the potential contribution of habitat heterogeneity and dispersal limitation to the formation of spatial patterns of tree species. The results showed that the inhomogeneous Thomas process was the best fit model and described most of the species studied, suggesting that spatial patterns of tree species might be formed by the joint effects of habitat associations and dispersal limitation. The homogeneous Thomas process that models the effect of dispersal limitation was the second best model. We also found that the best fit models could be predicted by species attributes, including species abundance and dispersal mode. The significant traits, however, differed between the two study plots and demonstrated site-specific patterns. This study indicated that the interactive operation of niche-based (habitat heterogeneity) and neutral-based (dispersal limitation) may be important in generating spatial patterns of tree species in forest communities.     

Tree Diversity,Forest Structure and Productivity along Altitudinal and Topographical Gradients in a Species-Rich Ecuadorian Montane Rain Forest

We studied the spatial heterogeneity of tree diversity, and of forest structure and productivity in a highly diverse tropical mountain area in southern Ecuador with the aim of understanding the causes of the large variation in these parameters. Two major environmental gradients, elevation and topography, representing a broad range of climatic and edaphic site conditions, were analyzed. We found the highest species richness of trees in valleys <2100 m. Valleys showed highest values of basal area, leaf area index and tree basal area increment as well. Tree diversity also increased from ridges to valleys, while canopy openness decreased. Significant relationships existed between tree diversity and soil parameters (pH, total contents of Mg, K, Ca, N and P), and between diversity and the spatial variability of pH and Ca and Mg contents suggesting a dependence of tree diversity on both absolute levels and on the small-scale heterogeneity of soil nutrient availability. Tree diversity and basal area increment were positively correlated, partly because both are similarly affected by soil conditions. We conclude that the extraordinarily high tree species richness in the area is primarily caused by three factors: (1) the existence of steep altitudinal and topographic gradients in a rather limited area creating a small-scale mosaic of edaphically different habitats; (2) the intermingling of Amazonian lowland plant species, that reach their upper distribution limits, and of montane forest species; and (3) the geographical position of the study area between the humid eastern Andean slope and the dry interandean forests of South Ecuador.     

Microbial biomass C and N,and mineralizable-N,in litter and mineral soil under Pinus radiata on a coastal sand: Influence of stand age and harvest management

Microbial biomass C and N, and anaerobically mineralizable-N, were measured in the litter and mineral soil (0–10 cm and 10–20 cm depth) of Pinus radiata plantations in two trials on a nitrogen-deficient coastal sand. The trials comprised (a) stands of different age (1 to 33 years), with five of the seven stands studied being second rotation, and (b) a harvest-management trial, with stands established after different harvesting treatments of the first-rotation trees and understorey development controlled by manual weeding and chemical sprays. The harvest-management stands were sampled in the fifth year after the second-rotation establishment.In the stands of different age, the levels of microbial biomass C and N, and also mineralizable-N, in the litter and mineral soil showed no relationship with tree age and were similar to those in the oldest (33 years) stands of P. radiata. In the harvesting trial, five years after establishment of the second rotation, levels of microbial N and mineralizable-N in the litter and mineral soil were generally lowest where whole trees and the original forest floor had been removed; they were higher in associated plots in which the original forest floor had been removed but fertilizer N was regularly applied. No marked differences were then found between the other harvest treatments, viz. whole-tree harvest, stem-only harvest with slash remaining on site, and stem-only harvest plus extra added slash materials. In each trial, levels of microbial C and N and mineralizable-N were closely related to total C, and especially total N, in 0–10 cm depth mineral soil, but not generally in litter. Respiratory measurements strongly suggest that the microbial populations in mineral soil had a high metabolic activity.On an area basis in the harvest-management trial, total tree N and microbial N in the litter and mineral soil were lowest in stands where the original forest floor had been removed. In this particular treatment, microbial N in the litter plus mineral soil (0–20 cm depth) after five years of second-rotation growth comprised 7.3% of the total ecosystem N; values in the other treatments ranged between 5.6 and 6.0%.Our results emphasise the importance of slash and litter, and probably volunteer shrubs and herbaceous under-storey species, in conserving pools of potentially available N during the early stages of tree development.     

Is Tree Species Diversity or Species Identity the More Important Driver of Soil Carbon Stocks,C/N Ratio,and pH?

We explored tree species diversity effects on soil C stock, C/N ratio, and pH as compared with effects of tree species identity. We sampled forest floors and mineral soil (0–40 cm) in a diversity gradient of 1–5 tree species composed of conifers and broadleaves in Bia?owie?a Forest, Poland. Diversity was a weaker driver than identity of soil C stocks, C/N ratio, and pH in the soil profile. However, there were significant non-additive effects of diversity and significant effects of identity on C stock and C/N ratio within different parts of the soil profile. More diverse forests had higher C stocks and C/N ratios in the 20–40 cm layer, whereas identity in terms of conifer proportion increased C stocks and C/N ratios only in forest floors. A positive relationship between C stocks and root biomass in the 30–40 cm layer suggested that belowground niche complementarity could be a driving mechanism for higher root carbon input and in turn a deeper distribution of C in diverse forests. Diversity and identity affected soil pH in topsoil with positive and negative impacts, respectively. More diverse forests would lead to higher soil nutrient status as reflected by higher topsoil pH, but there was a slight negative effect on N status as indicated by higher C/N ratios in the deeper layers. We conclude that tree species diversity increases soil C stocks and nutrient status to some extent, but tree species identity is a stronger driver of the studied soil properties, particularly in the topsoil.