The relationship between species richness and biomass changes from boreal to subtropical forests in China

Diversity‐manipulation experiments suggest a positive effect of biodiversity on ecosystem properties (EPs), but variable relationships between species richness and EPs have been reported in natural communities. An explanation for this discrepancy is that observed richness–EPs relationships in natural communities change with environment and species functional identities. But how the relationships change along broad‐scale climatic gradients has rarely been examined. In this paper, we sampled 848 plots of 20 × 30 m² from boreal to tropical forests across China. We examined plot biomass with respect to environmental factors, tree species richness and functional group identity (FGI, i.e. evergreen vs deciduous, and coniferous vs broadleaf). Variation partitioning was used to evaluate the relative effects of the three classes of factors. We found that, most of the ‘effects’ (percentage of variation explained) of richness and FGI on forest biomass were shared with environmental factors, but species richness and FGI still revealed significant effects in addition to environment for plots across China. Richness and FGI explained biomass mainly through their shared effects instead of independent effects, suggesting that the positive biodiversity effect is closely associated with a sampling effect. The relative effects of richness, FGI and environment varied latitudinally: the independent effects of environment and richness decreased from boreal to subtropical forests, whereas the total effect of FGI increased. We also found that the slope of richness–biomass relationship decreased monotonically from boreal to subtropical forests, possibly because of decreasing complementarity and increasing competition with increasing productivity. Our results suggest that while species richness does have significant effects on forest biomass it is less important than environmental gradients and other biotic factors in shaping large‐scale biomass patterns. We suggest that understanding how and why the diversity–EPs relationships change along climatic gradient would be helpful for a better understanding of real biodiversity effects in natural communities.     

Mass‐ratio and complementarity effects simultaneously drive aboveground biomass in temperate Quercus forests through stand structure

Forests play a key role in regulating the global carbon cycle, a substantial portion of which is stored in aboveground biomass (AGB). It is well understood that biodiversity can increase the biomass through complementarity and mass‐ratio effects, and the contribution of environmental factors and stand structure attributes to AGB was also observed. However, the relative influence of these factors in determining the AGB of Quercus forests remains poorly understood. Using a large dataset retrieved from 523 permanent forest inventory plots across Northeast China, we examined the effects of integrated multiple tree species diversity components (i.e., species richness, functional, and phylogenetic diversity), functional traits composition, environmental factors (climate and soil), stand age, and structure attributes (stand density, tree size diversity) on AGB based on structural equation models. We found that species richness and phylogenetic diversity both were not correlated with AGB. However, functional diversity positively affected AGB via an indirect effect in line with the complementarity effect. Moreover, the community‐weighted mean of specific leaf area and height increased AGB directly and indirectly, respectively; demonstrating the mass‐ratio effect. Furthermore, stand age, density, and tree size diversity were more important modulators of AGB than biodiversity. Our study highlights that biodiversity–AGB interaction is dependent on the regulation of stand structure that can be even more important for maintaining high biomass than biodiversity in temperate Quercus forests.     

广西马尾松林植物功能多样性与生产力的关系

探索植物多样性与生产力的关系可为森林经营与管理提供科学基础。本研究以广西4个地区的马尾松(Pinus massoniana)人工林群落为研究对象, 通过计算物种多样性、功能多样性和功能优势值, 运用相关分析、自动线性建模和方差划分等方法, 分析了多样性与生产力的关系。研究发现, 生产力与物种丰富度、Shannon指数、功能丰富度、功能均匀度极显著正相关(P < 0.01), 与物种均匀度、功能多样性、功能离散度、功能团个数、坡向显著正相关(P < 0.05), 与林龄极显著负相关(P < 0.01), 4个功能多样性参数和4个物种多样性参数两两之间皆为显著正相关; 未发现初始生物量制约生产力的提高; 方差划分最优模型中, 功能多样性参数效应、功能优势值效应和林龄效应分别解释生产力方差的56%、43%和33%, 功能多样性参数效应和功能优势值效应重叠部分高达27%; 生态位互补效应主要由功能丰富度和功能均匀度产生, 选择效应主要由生长型优势值产生; 生长型优势值为灌木的样地生产力较高, 次优种或过渡种对生态系统功能也有重要作用。以生产力为响应变量的自动线性建模最佳子集包括重要性由大到小的5个因素: 林龄、生长型优势值、功能丰富度、功能均匀度、功能团个数。建议维护森林功能多样性, 加强林下叶层植物保护, 用好功能重要的物种, 通过林下叶层的补偿性光合作用和生长竞争, 有效地提高生产力和生物多样性。     

Structural and taxonomic diversity predict above-ground biomass better than functional measures of maximum height in mixed-species forests

Aims: Mixed-species forests are known to be highly productive systems because of their high species diversity, including taxonomic diversity (species richness) and structural diversity. Recent empirical evidence also points to plant maximum height, as a functional trait that potentially drives forest above-ground biomass (AGB). However, the interrelations between these biotic variables are complex, and it is not always predictable if structural diversity attributes or functional metrics of plant maximum height would act as the most important determinant of stand biomass. Here we evaluated the relative importance of structural diversity attributes and functional metrics of plant maximum height (Hmax) in predicting and mediating AGB response to variation in species richness in mixed-species forests, while also accounting for fine-scale environmental variation. Location: Northern Benin. Methods: We used forest inventory data from mixed-species stands of native and exotic species. We quantified structural diversity as coefficient of variation of tree diameter at breast height (CVdbh) and of height (CVHt). For plant Hmax, we computed three metrics: functional range (FRHmax), functional divergence (FDHmax) and community-weighted mean (CWMHmax). We used topographical variables such as elevation and slope to account for possible environmental effects. Simple and multiple mixed-effects models, and structural equation models were performed to assess the direct and indirect links of AGB with species richness through structural diversity attributes and functional metrics of plant Hmax. Results: Species richness and CVdbh were positively related to AGB, while functional metrics of plant Hmax were not. Structural equation models revealed that species richness influenced AGB indirectly via CVdbh, which alone strongly promoted AGB. Elevation only had a positive direct effect on AGB. While increasing species richness enhanced CVdbh and functional measures of plant Hmax, there was no support for the latter mediating the effects of species richness on AGB. Conclusion: Structural diversity has a significant advantage in predicting and mediating the positive effect of species richness on AGB more so than functional measures of plant Hmax. We argue that structural diversity acts as a mechanism for the species richness–AGB relationship, and that maintaining high structural diversity would enhance biomass in mixed-species forests.     

Biodiversity and climate determine the functioning of Neotropical forests

Aim

Tropical forests account for a quarter of the global carbon storage and a third of the terrestrial productivity. Few studies have teased apart the relative importance of environmental factors and forest attributes for ecosystem functioning, especially for the tropics. This study aims to relate aboveground biomass (AGB) and biomass dynamics (i.e., net biomass productivity and its underlying demographic drivers: biomass recruitment, growth and mortality) to forest attributes (tree diversity, community‐mean traits and stand basal area) and environmental conditions (water availability, soil fertility and disturbance).

Location

Neotropics.

Methods

We used data from 26 sites, 201 1‐ha plots and >92,000 trees distributed across the Neotropics. We quantified for each site water availability and soil total exchangeable bases and for each plot three key community‐weighted mean functional traits that are important for biomass stocks and productivity. We used structural equation models to test the hypothesis that all drivers have independent, positive effects on biomass stocks and dynamics.

Results

Of the relationships analysed, vegetation attributes were more frequently associated significantly with biomass stocks and dynamics than environmental conditions (in 67 vs. 33% of the relationships). High climatic water availability increased biomass growth and stocks, light disturbance increased biomass growth, and soil bases had no effect. Rarefied tree species richness had consistent positive relationships with biomass stocks and dynamics, probably because of niche complementarity, but was not related to net biomass productivity. Community‐mean traits were good predictors of biomass stocks and dynamics.

Main conclusions

Water availability has a strong positive effect on biomass stocks and growth, and a future predicted increase in (atmospheric) drought might, therefore, potentially reduce carbon storage. Forest attributes, including species diversity and community‐weighted mean traits, have independent and important relationships with AGB stocks, dynamics and ecosystem functioning, not only in relatively simple temperate systems, but also in structurally complex hyper‐diverse tropical forests.     

Loss of biodiversity and shifts in aboveground biomass drivers in tropical rainforests with different disturbance histories

Tropical forests account for more than half of the global carbon forest stock and much of the biological diversity on Earth. However, disturbances such as deforestation and forest degradation threaten the maintenance of these ecosystem services. This study aimed to understand how different disturbance histories affect the forest stand biomass, as well as species and functional diversity, and to what extent these differences can change the relationships between biomass and their drivers. We used data from forests with clear-cut and selectively logged disturbance histories, and from old-growth forests, situated in the Brazilian Atlantic forest. Forests with logging disturbances showed significant losses in their aboveground biomass compared to those of old-growth forests (50% loss in selectively logged forests and 80% loss in clear-cut forests). Interestingly, only clear-cut secondary forests showed differences in species and functional diversity, and were dominated by species with acquisitive trait values, commonly found early in succession. Shifts in stand biomass drivers were observed in selectively logged forests. The mass-ratio hypothesis (mainly through the functional trait of maximum height) was the most important biomass driver in clear-cut secondary and old-growth forests, whereas the importance of the niche complementarity hypothesis (through functional richness and dispersion) was higher in selectively logged forests. Our study highlights that disturbance histories can affect forest aboveground biomass and its drivers. Moreover, our results reinforce the need for conservation of intact forests but highlight the importance of including degraded forests in conservation mechanisms based in carbon stocks, as these forests retain high values of species and functional diversities that are crucial to biomass and consequently carbon stock acquisition.     

Positive tree diversity effect on fine root biomass: via density dependence rather than spatial root partitioning

The importance of species richness to ecosystem functioning and services is a central tenet of biological conservation. However, most of our theory and mechanistic understanding is based on diversity found aboveground. Our study sought to better understand the relationship between diversity and belowground function by studying root biomass across a plant diversity gradient. We collected soil cores from 91 plots with between 1 and 12 aboveground tree species in three natural secondary forests to measure fine root (≤ 2 mm in diameter) biomass. Molecular methods were used to identify the tree species of fine roots and to estimate fine root biomass for each species. This study tested whether the spatial root partitioning (species differ by belowground territory) and symmetric growth (the capacity to colonize nutrient-rich hotspots) underpin the relationship between aboveground species richness and fine root biomass. All species preferred to grow in nutrient-rich areas and symmetric growth could explain the positive relationship between aboveground species richness and fine root biomass. However, symmetric growth only appeared in the nutrient-rich upper soil layer (0–10 cm). Structural equation modelling indicated that aboveground species richness and stand density significantly affected fine root biomass. Specifically, fine root biomass depended on the interaction between aboveground species richness and stand density, with fine root biomass increasing with species richness at lower stand density, but not at higher stand density. Overall, evidence for spatial (i.e. vertical) root partitioning was inconsistent; assumingly any roots growing into deeper unexplored soil layers were not sufficient contributors to the positive diversity–function relationship. Alternatively, density-dependent biotic interactions affecting tree recruitment are an important driver affecting productivity in diverse subtropical forests but the usual root distribution patterns in line with the spatial root partitioning hypothesis are unrealistic in contexts where soil nutrients are heterogeneously distributed.     

Recovery of mammal diversity in tropical forests: a functional approach to measuring restoration

Ecological restoration is increasingly applied in tropical forests to mitigate biodiversity loss and recover ecosystem functions. In restoration ecology, functional richness, rather than species richness, often determines community assembly, and measures of functional diversity provide a mechanistic link between diversity and ecological functioning of restored habitat. Vertebrate animals are important for ecosystem functioning. Here, we examine the functional diversity of small‐to‐medium sized mammals to evaluate the diversity and functional recovery of tropical rainforest. We assess how mammal species diversity and composition and functional diversity and composition, vary along a restoration chronosequence from degraded pasture to “old‐growth” tropical rainforest in the Wet Tropics of Australia. Species richness, diversity, evenness, and abundance did not vary, but total mammal biomass and mean species body mass increased with restoration age. Species composition in restoration forests converged on the composition of old‐growth rainforest and diverged from pasture with increasing restoration age. Functional metrics provided a clearer pattern of recovery than traditional species metrics, with most functional metrics significantly increasing with restoration age when taxonomic‐based metrics did not. Functional evenness and dispersion increased significantly with restoration age, suggesting that niche complementarity enhances species' abundances in restored sites. The change in community composition represented a functional shift from invasive, herbivorous, terrestrial habitat generalists and open environment specialists in pasture and young restoration sites, to predominantly endemic, folivorous, arboreal, and fossorial forest species in older restoration sites. This shift has positive implications for conservation and demonstrates the potential of tropical forest restoration to recover rainforest‐like, diverse faunal communities.     

氮添加对宁夏荒漠草原植物初级生产力的影响机制

许多研究探索了与全球变化相关的生态系统功能的变化,但对生态系统功能变化的机制与途径了解较少。初级生产力是生态系统功能的重要组分,但关于氮(N)添加下荒漠草原植物群落初级生产力如何变化以及变化机制尚未明确,N是否通过影响生物多样性来影响荒漠草原初级生产力?为此,本研究在荒漠草原开展了为期4年的N添加控制实验(2018—2021年),试验处理包括对照和4个N添加水平(5、10、20和40 g m^-2 a^-1),研究了N添加对荒漠草原物种多样性、功能多样性、初级生产力及其关系的影响。结果表明：(1)N添加处理(2018—2021年)改变了植物物种多样性及功能多样性,但年际间变化趋势不同。N添加处理第四年(2021年)荒漠草原植物功能多样性(Rao指数)、群落加权平均值-株高、功能均匀度和功能离散度均显著增加,而荒漠草原植物物种丰富度和Shannon-Wiener指数均显著降低。(2)N添加可以通过影响物种丰富度和功能多样性进而间接地促进荒漠草原初级生产力,但群落加权性状值-株高对初级生产力的影响是正效应,而物种丰富度和功能离散度对初级生产力的影响是...     

Tree diversity drives diversity of arthropod herbivores,but successional stage mediates detritivores

The high tree diversity of subtropical forests is linked to the biodiversity of other trophic levels. Disentangling the effects of tree species richness and composition, forest age, and stand structure on higher trophic levels in a forest landscape is important for understanding the factors that promote biodiversity and ecosystem functioning. Using a plot network spanning gradients of tree diversity and secondary succession in subtropical forest, we tested the effects of tree community characteristics (species richness and composition) and forest succession (stand age) on arthropod community characteristics (morphotype diversity, abundance and composition) of four arthropod functional groups. We posit that these gradients differentially affect the arthropod functional groups, which mediates the diversity, composition, and abundance of arthropods in subtropical forests. We found that herbivore richness was positively related to tree species richness. Furthermore, the composition of herbivore communities was associated with tree species composition. In contrast, detritivore richness and composition was associated with stand age instead of tree diversity. Predator and pollinator richness and abundance were not strongly related to either gradient, although positive trends with tree species richness were found for predators. The weaker effect of tree diversity on predators suggests a cascading diversity effect from trees to herbivores to predators. Our results suggest that arthropod diversity in a subtropical forest reflects the net outcome of complex interactions among variables associated with tree diversity and stand age. Despite this complexity, there are clear linkages between the overall richness and composition of tree and arthropod communities, in particular herbivores, demonstrating that these trophic levels directly impact each other.     

Water availability drives aboveground biomass and bird richness in forest restoration plantings to achieve carbon and biodiversity cobenefits

To combat global warming and biodiversity loss, we require effective forest restoration that encourages recovery of species diversity and ecosystem function to deliver essential ecosystem services, such as biomass accumulation. Further, understanding how and where to undertake restoration to achieve carbon sequestration and biodiversity conservation would provide an opportunity to finance ecosystem restoration under carbon markets. We surveyed 30 native mixed‐species plantings in subtropical forests and woodlands in Australia and used structural equation modeling to determine vegetation, soil, and climate variables most likely driving aboveground biomass accrual and bird richness and investigate the relationships between plant diversity, aboveground biomass accrual, and bird diversity. We focussed on woodland and forest‐dependent birds, and functional groups at risk of decline (insectivorous, understorey‐nesting, and small‐bodied birds). We found that mean moisture availability strongly limits aboveground biomass accrual and bird richness in restoration plantings, indicating potential synergies in choosing sites for carbon and biodiversity purposes. Counter to theory, woody plant richness was a poor direct predictor of aboveground biomass accrual, but was indirectly related via significant, positive effects of stand density. We also found no direct relationship between aboveground biomass accrual and bird richness, likely because of the strong effects of moisture availability on both variables. Instead, moisture availability and patch size strongly and positively influenced the richness of woodland and forest‐dependent birds. For understorey‐nesting birds, however, shrub cover and patch size predicted richness. Stand age or area of native vegetation surrounding the patch did not influence bird richness. Our results suggest that in subtropical biomes, planting larger patches to higher densities, ideally using a diversity of trees and shrubs (characteristics of ecological plantings) in more mesic locations will enhance the provision of carbon and biodiversity cobenefits. Further, ecological plantings will aid the rapid recovery of woodland and forest bird richness, with comparable aboveground biomass accrual to less diverse forestry plantations.     

Dispersal traits determine passive restoration trajectory of a Nigerian montane forest

Passive restoration methods offer great promise for tropical regions where resources are limited but the success of such efforts can be variable. Using trait-based theory, we investigated the likely trajectories of passive restoration efforts in a degraded Nigerian montane forest system recently protected from burning and cattle grazing. We quantified the density, species richness, and functional trait dispersion of dispersed seeds and seedling communities at increasing distances from the forest edge. We then determined which plant traits are responsible for colonisation by quantifying changes in functional-trait dispersion and relative frequencies of dispersal-linked traits with increasing distance from the forest. We found a rapid decrease in density and species richness, and significant species turnover in both seeds and seedlings just beyond the forest edge. This was mirrored by a significant decline in functional-trait dispersion and a shift in the relative frequencies of dispersal-linked traits. These findings suggest that the reassembly of plant communities adjacent to remnant forest is dependent on functional traits present in these remnant source populations, providing support for the incorporation of trait-based theory in restoration management.     

Biodiversity Promotes Tree Growth during Succession in Subtropical Forest

Losses of plant species diversity can affect ecosystem functioning, with decreased primary productivity being the most frequently reported effect in experimental plant assemblages, including tree plantations. Less is known about the role of biodiversity in natural ecosystems, including forests, despite their importance for global biogeochemical cycling and climate. In general, experimental manipulations of tree diversity will take decades to yield final results. To date, biodiversity effects in natural forests therefore have only been reported from sample surveys or meta-analyses with plots not initially selected for diversity. We studied biomass and growth of subtropical forests stands in southeastern China. Taking advantage of variation in species recruitment during secondary succession, we adopted a comparative study design selecting forest plots to span a gradient in species richness. We repeatedly censored the stem diameter of two tree size cohorts, comprising 93 species belonging to 57 genera and 33 families. Tree size and growth were analyzed in dependence of species richness, the functional diversity of growth-related traits, and phylogenetic diversity, using both general linear and structural equation modeling. Successional age covaried with diversity, but differently so in the two size cohorts. Plot-level stem basal area and growth were positively related with species richness, while growth was negatively related to successional age. The productivity increase in species-rich, functionally and phylogenetically diverse plots was driven by both larger mean sizes and larger numbers of trees. The biodiversity effects we report exceed those from experimental studies, sample surveys and meta-analyses, suggesting that subtropical tree diversity is an important driver of forest productivity and re-growth after disturbance that supports the provision of ecological services by these ecosystems.     

The species richness–productivity relationship in the herb layer of European deciduous forests

Aim In contrast to non‐forest vegetation, the species richness–productivity (SR‐P) relationship in forests still remains insufficiently explored. Several studies have focused on the diversity of the tree layer, but the species richness of temperate deciduous forests is mainly determined by their species‐rich herb layer. The factors controlling herb‐layer productivity may differ from those affecting tree layers or open herbaceous vegetation, and thus the SR‐P relationship and its underlying processes may differ. However, the few relevant studies have reported controversial results. Here we explore the SR‐P relationship in the forest herb layer across different areas from oceanic to continental Europe, and put the effect of habitat productivity on species richness into context with other key factors, namely soil pH and light availability. Location North‐western Germany, Czech Republic, Slovakia and southern Urals (Russia). Methods We measured herb‐layer species richness and biomass, soil pH and tree‐layer cover in 156 vegetation plots of 100 m² in deciduous forests. We analysed the SR‐P relationship and the relative importance of environmental variables using regression models for particular areas and separate forest types. Results We found a consistent monotonic increase in the herb‐layer species richness with productivity across all study areas and all forest types. Soil pH and light availability also affected species richness, but their relative importance differed among areas. Main conclusions We suggest that the monotonically increasing SR‐P relationship in the forest herb layer results from the fact that herb‐layer productivity is limited by canopy shading; competition within the herb layer is therefore not strong enough to exclude many species. This differs fundamentally from open herbaceous vegetation, which is not subject to such productivity limits and consequently exhibits a unimodal SR‐P relationship. We present a conceptual model that might explain the differences in the SR‐P relationship between the forest herb layer and open herbaceous vegetation.     

南亚热带常绿阔叶林树木多样性与生物量和生产力的关联及其影响因素

生物多样性和生态系统功能的关系直接或间接地影响着生产力, 是生态学研究的关键问题。本研究旨在定量探讨亚热带自然林演替后期森林生态系统树木多样性与生物量或生产力的关系。本研究基于中国南亚热带长期永久性样地的群落调查数据以及地形和土壤养分数据, 分析了南亚热带常绿阔叶林树木多样性与生物量和生产力的关联及其影响因素。相关性分析结果表明, 物种多样性与生物量呈显著负相关, 与生产力呈显著正相关; 结构多样性与生物量呈显著正相关, 与生产力呈显著负相关。此外, 不同环境因子对多样性、生物量和生产力的影响具有显著差异, 其中土壤含水量对生产力有显著影响, 物种多样性指标与部分地形和土壤因子均有相关性, 而群落结构多样性指标与土壤因子的相关性更强。方差分解结果表明, 结构多样性对生物量和生产力的单独效应的解释率最大, 分别为35.39%和5.21%; 其次是结构多样性和物种多样性的共同效应, 对生物量和生产力的解释率分别为13.66%和3.53%; 地形和土壤因子的解释率较小。同时, 结构方程结果也表明, 结构多样性对生物量有较强的直接正影响; 生物量对生产力有强烈的直接负影响, 结构多样性通过增加生物量明显地减少了生产力; 土壤和地形因子主要是通过物种和结构多样性间接影响生物量和生产力。综上, 本研究认为在南亚热带森林演替顶极群落中, 群落结构复杂性和物种多样性的提高对促进群落生产力和生物量具有重要作用。     

Effect of forest landscapes composition and configuration on bird community and its functional traits in a hotspot of biodiversity of Chile

Understanding the effect of landscape configuration on the bird species richness and their functional traits (dietary preferences) is important to link the conservation and restoration plans to the production of the crops. Our aims were: 1) to study the influence of forest types (native, mixed and plantations) on the bird species richness in two agroforestry landscapes (heterogeneous/homogeneous); 2) to assess the effect of size/density of forest patches in the birds’ functional traits; 3) to evaluate the effect of isolated trees on them, and 4) to discuss conservation and restoration measures for the birds’ functional traits in agroforestry landscapes. We used hierarchical occupancy models to evaluate the effect of different landscape metrics and detectability measures on bird communities. We recorded a total of 64 bird species. The estimated species richness was considerable higher in homogeneous landscape (31.7 ± 2.7) than heterogeneous (27.3 ± 2.5). Our results showed the bird assemblage had a positive trend with native forests, negative with mixed forests and neutral trend for plantations. The granivores and insectivore’s species showed significant preferences for homogeneous landscape, while omnivores had significant preferences for heterogeneous landscape. Carnivores/Piscivores and herbivores/frugivores species did not show preferences by any landscape type. The response of functional traits depended on different forests attributes. The isolated trees had a significant effect on the birds’ functional traits. In conclusion, it is necessary a deep knowledge about the relationship between the landscape configuration and the bird species richness/functional traits. These findings could help in the future with the conservation, restoration, and rewilding policies in this important hotspot of biodiversity, avoiding alterations in the ecosystem services.     

Dominance by an obligate annual affects the morphological characteristics and biomass production of a planted wetland macrophyte community

Aims Biodiversity–ecosystem function experiments can test for causal relationships between planting diversity and community productivity. Planting diversity is routinely introduced as a design element in created wetlands, yet substantive support for the finding that early diversity positively affects ecosystem functioning is lacking for wetlands. We conducted a 2-year diversity–productivity experiment using freshwater wetland mesocosms to investigate community biomass production as affected by planted macrophyte functional richness.Methods A richness gradient of macrophytes in four emergent wetland plant functional groups was established in freshwater mesocosms for two consecutive years. Species-specific aboveground morphological traits of plant size were measured at peak growth in both years; rooting depth was measured for each species in the second year. Aboveground biomass (AGB) and belowground biomass (BGB) were harvested after peak growth in the second year; first year AGB was estimated from morphological traits in constructed regression equations. Net richness effects (i.e. both complementarity effects and selection effects) were calculated using an additive partitioning method.Important findings Species richness had a positive effect on community AGB relative to monocultures in the first year. In the second year, mean AGB was significantly reduced by competition in the most species-rich mixtures and all mixtures underyielded relative to the average monoculture. Competition for soil resources was weaker belowground, whereby root distribution at depths>20cm was reduced at the highest richness levels but overall BGB production was not affected. Changes in species biomass were strongly reflected by variation in species morphological traits, and species above and belowground performances were highly correlated. The obligate annual (Eleocharis obtusa), a dominant competitor, significantly contributed to the depression of perennial species' growth in the second growing season. To foster primary productivity with macrophyte richness in early successional communities of created wetlands where ruderal strategies are favored and competition may be stronger than species complementarity, unsystematic planting designs such as clustering the same or similar species could provide protection for some individuals. Additionally, engineering design elements fostering spatial or temporal environmental variability (e.g. microtopography) in newly created wetlands helps diversify the responses of wetland macrophyte species to their environment and could allow for greater complementarity in biomass production.     

Plant species diversity in Mediterranean old-growth forests: A case study from central Italy

Abstract

To investigate the differences in understorey composition and diversity between old-growth and managed forests, we analyzed an old-growth and a managed beech stand in the same area displaying similar abiotic features. We considered variations in understorey species composition and richness. The sampled understorey species were characterized in terms of functional traits, Ellenberg's indicator values and taxonomic distinctness; next, we calculated four different pairwise plot-to-plot dissimilarity matrices based on species composition, functional traits, Ellenberg's indices and taxonomic distances. We applied a permutational multivariate extension of ANOVA to test whether the forest stands significantly differ in the considered features. Indicator values of all plant species in managed and old-growth stands were evaluated.

The old-growth forest had a higher species richness; permutational analysis of variance showed significant differences between the two stands in plant species composition, functional traits, Ellenberg indices and taxonomic distances. Indicator species analysis highlighted 14 indicator species for the unmanaged stand, while only 3 indicators were found for the managed one.

The results suggest that forest management determines ecological differences that strongly affect plant species composition.

The knowledge of natural stands dynamics could allow development of new approaches and practices in forest management focusing on biodiversity conservation.     

Comparing the recovery of richness,structure, and biomass in naturally regrowing and planted reforestation

The clearing of natural vegetation for agriculture has reduced the capacity of natural systems to provide ecosystem functions. Ecological restoration can restore desirable ecosystem functions, such as creating habitat for animal conservation and carbon sequestration as woody biomass. In order to maintain these beneficial ecosystem functions, restoration projects need to mature into self‐perpetuating communities. Here we compared the ecological attributes of two types of restoration, “active” tree plantings with “passive” natural forest regeneration (“natural regrowth”) to existing remnant vegetation in a cleared agricultural landscape. Specifically, we measured differences between forest categories in factors that may predict future restoration failure or ecosystem collapse: aboveground plant biomass and biomass accrual over time (for regrowing stands), plant density and size class distributions, and diversity of functional groups based on seed dispersal and growth strategy traits. We found that natural regrowth and planted forests were similar in many ecological characteristics, including biomass accrual. Despite this, planted stands contained fewer tree recruit and shrub individuals, which may be due to limited recruitment in plantings. If this continues, these forests may be at risk of collapsing into nonforest states after mature trees senesce. Lower shrub density and richness of mid‐story trees may lead to lower structural complexity in planting plots, and alongside lower richness of fleshy‐fruited plant species may reduce animal resources and animal use of the restored stand. In our study region, natural regrowth may result in restored woodland communities with greater conservation and carbon mitigation value.     

Functional traits determine tree growth and ecosystem productivity of a tropical montane forest: Insights from a long‐term nutrient manipulation experiment

Trait‐response effects are critical to forecast community structure and biomass production in highly diverse tropical forests. Ecological theory and few observation studies indicate that trees with acquisitive functional traits would respond more strongly to higher resource availability than those with conservative traits. We assessed how long‐term tree growth in experimental nutrient addition plots (N, P, and N + P) varied as a function of morphological traits, tree size, and species identity. We also evaluated how trait‐based responses affected stand scale biomass production considering the community structure. We found that tree growth depended on interactions between functional traits and the type or combination of nutrients added. Common species with acquisitive functional traits responded more strongly to nutrient addition, mainly to N + P. Phosphorous enhanced the growth rates of species with acquisitive and conservative traits, had mostly positive effects on common species and neutral or negative effects in rare species. Moreover, trees receiving N + P grew faster irrespective of their initial size relative to trees in control or to trees in other treatment plots. Finally, species responses were highly idiosyncratic suggesting that community processes including competition and niche dimensionality may be altered under increased resource availability. We found no statistically significant effects of nutrient additions on aboveground biomass productivity because acquisitive species had a limited potential to increase their biomass, possibly due to their generally lower wood density. In contrast, P addition increased the growth rates of species characterized by more conservative resource strategies (with higher wood density) that were poorly represented in the plant community. We provide the first long‐term experimental evidence that trait‐based responses, community structure, and community processes modulate the effects of increased nutrient availability on biomass productivity in a tropical forest.