Scale‐dependent effects of neighborhood biodiversity on individual tree productivity in a coniferous and broad‐leaved mixed forest in China

The relationship between biodiversity and productivity has stimulated an increasing body of research over the past decades, and this topic still occupies a central place in ecology. While most studies have focused on biomass production in quadrats or plots, few have investigated the scale‐dependent relationship from an individual plant perspective. We present an analysis of the effects of biodiversity (species diversity and functional diversity) on individual tree growth with a data set of 16,060 growth records from a 30‐ha temperate forest plot using spatially explicit individual tree‐based methods. A significant relationship between species diversity and tree growth was found at the individual tree level in our study. The magnitude and direction of biodiversity effects varies with the spatial scale. We found positive effects of species diversity on tree growth at scales exceeding 9 m. Individual tree growth rates increased when there was a greater diversity of species in the neighborhood of the focal tree, which provides evidence of a niche complementarity effect. At small scales (3–5 m), species diversity had negative effects on tree growth, suggesting that competition is more prevalent than complementarity or facilitation in these close neighborhoods. The results also revealed many confounding factors which influence tree growth, such as elevation and available sun light. We conclude that the use of individual tree‐based methods may lead to a better understanding of the biodiversity‐productivity relationship in forest communities.     

A role for the sampling effect in invaded ecosystems

Species loss and invasion of exotic species are two components of global biodiversity change that are expected to influence ecosystem functioning. Yet how they interact in natural settings remains unclear. Experiments have revealed two major mechanisms for the observed increase in primary productivity with plant species richness. Plant productivity may rise with species richness due to the increased amount of resources used by more diverse communities (niche complementarity) or through the increased probability of including a highly productive, dominant species in the community (sampling effect). Current evidence suggests that niche complementarity is the most relevant mechanism, whereas the sampling effect would only play a minor and transient role in natural systems. In turn, exotic species can invade by using untapped resources or because they possess a fitness advantage over resident species allowing them to dominate the community. We argue that the sampling effect can be a significant biodiversity mechanism in ecosystems invaded by dominant exotic species, and that the effect can be persistent even after decades of succession. We illustrate this idea by analyzing tree species richness–productivity relationships in a subtropical montane forest (NW Argentina) heavily invaded by Ligustrum lucidum, an evergreen tree from Asia. We found that the forest biomass increased along a natural gradient of tree species richness whether invaded by L. lucidum or not. Consistent with the sampling effect, L. lucidum invasion tripled total tree biomass irrespective of species richness, and monocultures of L. lucidum were more productive than any of the most species‐rich, uninvaded communities. Hence, the sampling effect may not be restricted to randomly assembled, synthetic communities. We emphasize that studying invaded ecosystems may provide novel insights on the mechanisms underlying the effect of biodiversity on ecosystem function.     

The effect of biodiversity on tree productivity: from temperate to boreal forests

Aim An important issue regarding biodiversity concerns its influence on ecosystem functioning. Experimental work has led to the proposal of mechanisms such as niche complementarity. However, few attempts have been made to confirm these in natural systems, especially in forests. Furthermore, one of the most interesting unresolved questions is whether the effects of complementarity on ecosystem functioning (EF) decrease in favour of competitive exclusions over an increasing productivity gradient. Using records from permanent forest plots, we asked the following questions. (1) Is tree productivity positively related to diversity? (2) Does the effect of diversity increase in less productive forests? (3) What metric of diversity (e.g. functional or phylogenetic diversity) better relates to tree productivity? Location Temperate, mixed and boreal forests of eastern Canada. Methods Over 12,000 permanent forest plots, from temperate to boreal forests, were used to test our hypotheses in two steps. (1) Stepwise regressions were used to identify the best explanatory variables for tree productivity. (2) The selected climatic and environmental variables, as well as density and biodiversity indices, were included in a structural equation model where links (paths) between covarying variables are made explicit, making structural equation modelling the best tool to explore such complicated causal networks. Results This is the first large‐scale demonstration of a strong, positive and significant effect of biodiversity on tree productivity with control for climatic and environmental conditions. Important differences were noted between the two forest biomes investigated. Main conclusions We show for the first time that complementarity may be less important in temperate forests growing in a more stable and productive environment where competitive exclusion is the most probable outcome of species interactions, whereas in the more stressful environment of boreal forests, beneficial interactions between species may be more important. The present work is also a framework for the analysis of large datasets in biodiversity–ecosystem functioning (B‐EF) research.     

Integrating mechanistic and empirical model projections to assess climate impacts on tree species distributions in northwestern North America

Empirical and mechanistic models have both been used to assess the potential impacts of climate change on species distributions, and each modeling approach has its strengths and weaknesses. Here, we demonstrate an approach to projecting climate‐driven changes in species distributions that draws on both empirical and mechanistic models. We combined projections from a dynamic global vegetation model (DGVM) that simulates the distributions of biomes based on basic plant functional types with projections from empirical climatic niche models for six tree species in northwestern North America. These integrated model outputs incorporate important biological processes, such as competition, physiological responses of plants to changes in atmospheric CO₂ concentrations, and fire, as well as what are likely to be species‐specific climatic constraints. We compared the integrated projections to projections from the empirical climatic niche models alone. Overall, our integrated model outputs projected a greater climate‐driven loss of potentially suitable environmental space than did the empirical climatic niche model outputs alone for the majority of modeled species. Our results also show that refining species distributions with DGVM outputs had large effects on the geographic locations of suitable habitat. We demonstrate one approach to integrating the outputs of mechanistic and empirical niche models to produce bioclimatic projections. But perhaps more importantly, our study reveals the potential for empirical climatic niche models to over‐predict suitable environmental space under future climatic conditions.     

Phylogenetic turnover in tropical tree communities: impact of environmental filtering,biogeography and mesoclimatic niche conservatism

Aim We addressed the roles of environmental filtering, historical biogeography and evolutionary niche conservatism on the phylogenetic structure of tropical tree communities with the following questions. (1) What is the impact of mesoclimatic gradients and dispersal limitation on phylogenetic turnover and species turnover? (2) How does phylogenetic turnover between continents compare in intensity with the turnover driven by climatic gradients at a regional scale? (3) Are independent phylogenetic reconstructions of the mesoclimatic niche of clades congruent between continents? Location Panama Canal Watershed and Western Ghats (India), two anciently divergent biogeographic contexts but with comparable rainfall gradients. Methods Using floristic data for 50 1‐ha plots in each region, independent measures of phylogenetic turnover (Π_ST) and species turnover (Jaccard) between plots were regressed on geographic and ecological distances. Mesoclimatic niches were reconstructed for each node of the phylogeny and compared between the two continents. Results (1) The phylogenetic turnover within each region is best explained by mesoclimatic differences (environmental filtering), while species turnover depends both on mesoclimatic differences and geographic distances (dispersal limitation). (2) The phylogenetic turnover between continents (Π_ST = 0.009) is comparable to that caused by mesoclimatic gradients within regions (Π_ST = 0.010) and both effects seem cumulative. (3) Independent phylogenetic reconstructions of the mesoclimatic niches were strongly correlated between the two continents (r = 0.61), despite the absence of shared species. Main conclusions Our results demonstrate a world‐wide deep phylogenetic signal for mesoclimatic niche within a biome, indicating that positive phylogenetic turnover at a regional scale reflects environmental filtering in plant communities.     

Macroecology, global change and the shadow of forgotten ancestors

Many recent studies have evaluated how global changes will affect biodiversity, and have mainly focused on how to develop conservation strategies to avoid, or at least minimize, extinctions due to shifts in suitable habitats for the species. However, these complex potential responses might be in part structured in phylogeny, because of the macroecological traits underlying them. In this comment, we review recent analytical developments in phylogenetic comparative methods that can be used to understand patterns of trait changes under environmental change. We focus on a partial regression approach that allows for partitioning the variance of traits into a fraction attributed to a pure ecological component, a fraction attributed to phylogenetically structured environmental variation (niche conservatism) and a fraction that may be attributed to phylogenetic effects only. We then develop a novel interpretation for linking these components for multiple traits with potential responses of species to global environmental change (i.e. adaptation, range shifts or extinctions). We hope that this interpretation will stimulate further research linking evolutionary components of multiple traits with broad-scale environmental changes.     

Experimental evaluation of diversity–productivity relationships in a coral reef fish assemblage

The global decline in biodiversity is causing increasing concern about the effects of biodiversity loss on ecosystem services such as productivity. Biodiversity has been hypothesised to be important in maintaining productivity of biological assemblages because niche complementarity and facilitation among the constituent species can result in more efficient use of resources. However, these conclusions are primarily based on studies with plant communities, and the relationship between diversity and productivity at higher trophic levels is largely unknown, especially in the marine environment. Here, we used a manipulative field experiment to test the effects of species richness and species identity on biomass accumulation in coral reef fish assemblages at Lizard Island. Small patch reefs were stocked with a total of 30 juveniles belonging to three planktivorous damselfish (genus Pomacentrus) according to three different levels of fish species richness (one, two and three species) and seven different combinations of fish species. Species richness had no effect on the relative growth in this assemblage after 18 days, but relative growth differed among individual fish species and the different combinations of species. Patterns of increase in biomass were best explained by species-specific differences and variable effects of intra- and interspecific competition on growth. These results suggest that niche complementarity and facilitation are not the most influential drivers of total productivity within this guild of planktivorous fishes. Total productivity may be resilient to declining reef fish biodiversity, but this will depend on which species are lost and on the life-history traits of remaining species.     

择伐对思茅松天然林乔木种间与种内关系的影响

通过分析不同择伐强度下思茅松(Pinus kesiya)天然林乔木种群的生态位、种间联结、种内与种间竞争和群落稳定性, 探讨干扰强度对思茅松天然林乔木种群种间与种内关系的影响。结果表明: (1)择伐干扰改变了大多数乔木树种在群落中的优势地位, 思茅松优势地位未受影响, 随着择伐干扰强度的增加, 思茅松在群落中的生态位宽度逐渐变大, 在重度干扰的群落中与红木荷(Schima wallichii)一起成为生态幅度最大的树种。(2)通过种间联结与生态位重叠分析发现, 随着干扰强度的加剧, 乔木种群的种间关系发生变化, 中度干扰群落中物种之间的生态位重叠明显高于未干扰和重度干扰群落, 而在三类群落中, 思茅松与其他种群之间的重叠值较小。随着干扰强度增加, 种群间正联结与负联结的种对所占的比例逐渐增加。(3) Hegyi竞争指数表明, 三类群落中思茅松种群的竞争压力主要来自种间, 而干扰强度进一步增加了思茅松种群的种间竞争强度, 其种内平均竞争指数随着对象木胸径的增加而减少。(4)林窗是思茅松种群种间和种内关系变化的主要原因, 三类群落都处于较不稳定的阶段, 但中度干扰群落的物种丰富度与稳定性都高于未干扰和重度干扰的群落。择伐原则依据森林管理的目的而确定, 如以木材培育为主或森林生态系统服务功能为主。     

From competition to facilitation: how tree species respond to neighbourhood diversity

Studies on tree communities have demonstrated that species diversity can enhance forest productivity, but the driving mechanisms at the local neighbourhood level remain poorly understood. Here, we use data from a large‐scale biodiversity experiment with 24 subtropical tree species to show that neighbourhood tree species richness generally promotes individual tree productivity. We found that the underlying mechanisms depend on a focal tree's functional traits: For species with a conservative resource‐use strategy diversity effects were brought about by facilitation, and for species with acquisitive traits by competitive reduction. Moreover, positive diversity effects were strongest under low competition intensity (quantified as the total basal area of neighbours) for acquisitive species, and under high competition intensity for conservative species. Our findings demonstrate that net biodiversity effects in tree communities can vary over small spatial scales, emphasising the need to consider variation in local neighbourhood interactions to better understand effects at the community level.     

Tree species richness promotes productivity in temperate forests through strong complementarity between species

Understanding the link between biodiversity and ecosystem functioning (BEF) is pivotal in the context of global biodiversity loss. Yet, long-term effects have been explored only weakly, especially for forests, and no clear evidence has been found regarding the underlying mechanisms. We explore the long-term relationship between diversity and productivity using a forest succession model. Extensive simulations show that tree species richness promotes productivity in European temperate forests across a large climatic gradient, mostly through strong complementarity between species. We show that this biodiversity effect emerges because increasing species richness promotes higher diversity in shade tolerance and growth ability, which results in forests responding faster to small-scale mortality events. Our study generalises results from short-term experiments in grasslands to forest ecosystems and demonstrates that competition for light alone induces a positive effect of biodiversity on productivity, thus providing a new angle for explaining BEF relationships.     

Parapatry and niche complementarity of Peruvian Desert geckos (Phyllodactylus): the ambiguous role of competition

The Sechura Desert of Peru is among the most arid, barren regions of South America. Four species of nocturnal geckos (Phyllodactylus) are parapatric in part of the desert. By comparing niche associations of these species in allopatry and parapatry, I attempt to determine whether the observed parapatric distributions and niche dimension complementarity are related to competition — as is frequently assumed. While parapatry suggests a role for competition, distributional patterns can alternatively be related to adaptations of geckos to different physical environments (sandy desert and rocky foothill) that abut in the study area. Niche complementarity might also be a result of competition, but potentially contradictory evidence suggests that niche complementarity might instead be the result of adaptations developed in allopatry and having no relationship to competition. The ambiguity of these interpretations sets limits on the significance of this kind of evidence: in the absence of attempts to falsify alternative explanations, observations of parapatry or of niche dimension complementarity do not demonstrate conclusively the impact of competition as a force structuring communities.     

Does species diversity limit productivity in natural grassland communities?

Theoretical analyses and experimental studies of synthesized assemblages indicate that under particular circumstances species diversity can enhance community productivity through niche complementarity. It remains unclear whether this process has important effects in mature natural ecosystems where competitive feedbacks and complex environmental influences affect diversity–productivity relationships. In this study, we evaluated diversity–productivity relationships while statistically controlling for environmental influences in 12 natural grassland ecosystems. Because diversity–productivity relationships are conspicuously nonlinear, we developed a nonlinear structural equation modeling (SEM) methodology to separate the effects of diversity on productivity from the effects of productivity on diversity. Meta-analysis was used to summarize the SEM findings across studies. While competitive effects were readily detected, enhancement of production by diversity was not. These results suggest that the influence of small-scale diversity on productivity in mature natural systems is a weak force, both in absolute terms and relative to the effects of other controls on productivity.     

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.     

Drivers of understorey biomass: tree species identity is more important than richness in a young forest

林下生物量影响因素：幼龄林树种特性比丰富度更重要 生物多样性与生态系统功能的正相关关系已被广泛报道,其主要来源于对草原生态系统的研究。然而,该结论并不一定适用于更复杂的环境,例如具有不同垂直层次的森林。举例而言,已有研究表明上层乔木树种丰富度与林下生产力降低有关。树种丰富度是否会通过增加(由于生境异质性)或降低(通过增强竞争)资源的可利用性进而影响林下生产力,以及林下生产力是否受树种特性的影响更大,这些影响机制都可能会随着时间的推移而改变。此外,研究还表明,丰富度-生产力关系随着环境背景的变化而改变。本研究利用可以操控树种丰富度的实验林场研究了这些不同垂直层位里的时间和环境动态。在中国亚热带森林生物多样性与生态系统功能(BEF-China)研究计划的框架下,我们在3年时间里沿树种丰富度梯度反复采集林下生物量样本,研究了不同环境处理中树种丰富度、树种特性和时间对林下生物量的影响。尽管我们发现乔木层特性对林下生物量有显著和一致的影响,但是树种丰富度对后者却不具有这种影响。另外,在森林结构层之间,可能并不存在单一的、具有普遍性的上层乔木树种丰富度与林下生产力的相关关系,并且与上层乔木相关的环境因素(如透光率)对林下生产力的贡献程度会随着时间而变化。总体而言,我们的结果表明,在研究森林结构层之间的关系时应将时间动态变化考虑在内。     

When do herbivorous insects compete? A phylogenetic meta‐analysis

When herbivorous insects interact, they can increase or decrease each other's fitness. As it stands, we know little of what causes this variation. Classic competition theory predicts that competition will increase with niche overlap and population density. And classic hypotheses of herbivorous insect diversification predict that diet specialists will be superior competitors to generalists. Here, we test these predictions using phylogenetic meta‐analysis. We estimate the effects of diet breadth, population density and proxies of niche overlap: phylogenetic relatedness, physical proximity and feeding‐guild membership. As predicted, we find that competition between herbivorous insects increases with population density as well as phylogenetic and physical proximity. Contrary to predictions, competition tends to be stronger between than within feeding guilds and affects specialists as much as generalists. This is the first statistical evidence that niche overlap increases competition between herbivorous insects. However, niche overlap is not everything; complex feeding guild effects indicate important indirect interactions.     

Deep phylogeny,net primary productivity,and global body size gradient in birds

Body size is evolutionarily constrained, but the influence of phylogenetic relationships on global body size (i.e. body mass) gradients is unexplored. We quantify and map the family‐level phylogenetic and non‐phylogenetic structure of the global gradient of birds, evaluating the extent to which it is influenced by phylogenetic inertia in contrast to heat conservation, resource availability, starvation resistance, niche conservatism, or interspecific competition. Phylogenetic eigenvector regression (PVR) partitioned the global bird body size gradient into phylogenetically autocorrelated (PA) and phylogenetically independent (PI) components. Simple, piecewise, and partial regressions were used to investigate associations between the PA and PI components of body size and environmental correlates, and to quantify independent and overlapping contributions of environment, phylogenetic autocorrelation, and species richness to the body size gradient. Two‐thirds of the geographic variation in bird body size can be explained by phylogenetic relationships at the family level. The global variation in body size, independent of phylogenetic relationships, is most strongly associated with net primary productivity, which is consistent with ‘starvation resistance’. However, the New and Old worlds have very different patterns. We found no independent association of species richness with body size. Despite major unresolved regional differences, deep phylogenetic relationships, heat conservation, and starvation resistance probably operate in concert in shaping the global bird body size gradient in different parts of the world. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

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.     

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.     

Conserving species‐rich predator assemblages strengthens natural pest control in a climate warming context

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Evolutionary History and Novel Biotic Interactions Determine Plant Responses to Elevated CO2 and Nitrogen Fertilization

A major frontier in global change research is predicting how multiple agents of global change will alter plant productivity, a critical component of the carbon cycle. Recent research has shown that plant responses to climate change are phylogenetically conserved such that species within some lineages are more productive than those within other lineages in changing environments. However, it remains unclear how phylogenetic patterns in plant responses to changing abiotic conditions may be altered by another agent of global change, the introduction of non-native species. Using a system of 28 native Tasmanian Eucalyptus species belonging to two subgenera, Symphyomyrtus and Eucalyptus, we hypothesized that productivity responses to abiotic agents of global change (elevated CO₂ and increased soil N) are unique to lineages, but that novel interactions with a non-native species mediate these responses. We tested this hypothesis by examining productivity of 1) native species monocultures and 2) mixtures of native species with an introduced hardwood plantation species, Eucalyptus nitens, to experimentally manipulated soil N and atmospheric CO₂. Consistent with past research, we found that N limits productivity overall, especially in elevated CO₂ conditions. However, monocultures of species within the Symphyomyrtus subgenus showed the strongest response to N (gained 127% more total biomass) in elevated CO₂ conditions, whereas those within the Eucalyptus subgenus did not respond to N. Root:shoot ratio (an indicator of resource use) was on average greater in species pairs containing Symphyomyrtus species, suggesting that functional traits important for resource uptake are phylogenetically conserved and explaining the phylogenetic pattern in plant response to changing environmental conditions. Yet, native species mixtures with E. nitens exhibited responses to CO₂ and N that differed from those of monocultures, supporting our hypothesis and highlighting that both plant evolutionary history and introduced species will shape community productivity in a changing world.