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.     

Drivers of productivity and its temporal stability in a tropical tree diversity experiment

There is increasing evidence that mixed‐species forests can provide multiple ecosystem services at a higher level than their monospecific counterparts. However, most studies concerning tree diversity and ecosystem functioning relationships use data from forest inventories (under noncontrolled conditions) or from very young plantation experiments. Here, we investigated temporal dynamics of diversity–productivity relationships and diversity–stability relationships in the oldest tropical tree diversity experiment. Sardinilla was established in Panama in 2001, with 22 plots that form a gradient in native tree species richness of one‐, two‐, three‐ and five‐species communities. Using annual data describing tree diameters and heights, we calculated basal area increment as the proxy of tree productivity. We combined tree neighbourhood‐ and community‐level analyses and tested the effects of both species diversity and structural diversity on productivity and its temporal stability. General patterns were consistent across both scales indicating that tree–tree interactions in neighbourhoods drive observed diversity effects. From 2006 to 2016, mean overyielding (higher productivity in mixtures than in monocultures) was 25%–30% in two‐ and three‐species mixtures and 50% in five‐species stands. Tree neighbourhood diversity enhanced community productivity but the effect of species diversity was stronger and increased over time, whereas the effect of structural diversity declined. Temporal stability of community productivity increased with species diversity via two principle mechanisms: asynchronous responses of species to environmental variability and overyielding. Overyielding in mixtures was highest during a strong El Niño‐related drought. Overall, positive diversity–productivity and diversity–stability relationships predominated, with the highest productivity and stability at the highest levels of diversity. These results provide new insights into mixing effects in diverse, tropical plantations and highlight the importance of analyses of temporal dynamics for our understanding of the complex relationships between diversity, productivity and stability. Under climate change, mixed‐species forests may provide both high levels and high stability of production.     

Nitrogen enrichment weakens ecosystem stability through decreased species asynchrony and population stability in a temperate grassland

Biodiversity generally promotes ecosystem stability. To assess whether the diversity–stability relationship observed under ambient nitrogen (N) conditions still holds under N enriched conditions, we designed a 6‐year field experiment to test whether the magnitude and frequency of N enrichment affects ecosystem stability and its relationship with species diversity in a temperate grassland. Results of this experiment showed that the frequency of N addition had no effect on either the temporal stability of ecosystem and population or the relationship between diversity and stability. Nitrogen addition decreased ecosystem stability significantly through decreases in species asynchrony and population stability. Species richness was positively associated with ecosystem stability, but no significant relationship between diversity and the residuals of ecosystem stability was detected after controlling for the effects of the magnitude of N addition, suggesting collinearity between the effects of N addition and species richness on ecosystem stability, with the former prevailing over the latter. Both population stability and the residuals of population stability after controlling for the effects of the magnitude of N addition were positively associated with ecosystem stability, indicating that the stabilizing effects of component populations were still present after N enrichment. Our study supports the theory predicting that the effects of environmental factors on ecosystem functioning are stronger than those of biodiversity. Understanding such mechanisms is important and urgent to protect biodiversity in mediating ecosystem functioning and services in the face of global changes.     

Differential roles of species richness versus species asynchrony in regulating community stability along a precipitation gradient

Plant community may provide products and services to humans. However, patterns and drivers of community stability along a precipitation gradient remain unclear. A regional‐scale transect survey was conducted over a 3‐year period from 2013 to 2015, along a precipitation gradient from 275 to 555 mm and spanning 440 km in length from west to east in a temperate semiarid grassland of northern China, a central part of the Eurasian steppe. Our study provided regional‐scale evidence that the community stability increased with increasing precipitation in the semiarid ecosystem. The patterns of community stability along a precipitation gradient were ascribed to community composition and community dynamics, such as species richness and species asynchrony, rather than the abiotic effect of precipitation. Species richness regulated the temporal mean (μ) of aboveground net primary productivity (ANPP), while species asynchrony regulated the temporal standard deviation (σ) of ANPP, which in turn contributed to community stability. Our findings highlight the crucial role of community composition and community dynamics in regulating community stability under climate change.     

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.     

物种多样性和系统发育多样性对阔叶红松林生产力的影响

生物多样性与生态系统功能间的关系已成为生态学研究的热点问题之一,其中植物多样性对森林生产力的驱动作用受到广泛关注,而其潜在驱动机制还存在很大争议.本研究依托黑龙江凉水国家级自然保护区典型阔叶红松林9 hm^2森林动态监测样地,利用2005年和2015年的调查数据,采用线性回归和结构方程模型探究不同空间尺度下物种多样性和系统发育多样性对森林生产力的影响.结果表明:物种多样性和系统发育多样性与生产力均呈正相关,随着空间尺度的增大,物种多样性对生产力的作用强度逐渐增强,而系统发育多样性对生产力的作用逐渐减弱;小尺度下系统发育多样性对生产力的影响大于物种多样性.生产力还受到非生物因素影响,在不同尺度下土壤因子与生产力均呈显著正相关,并且随着尺度的增大,土壤因子对生产力的作用逐渐占据主导地位.在今后研究中应将进化信息与生态系统功能相联系,可为其他多样性度量提供额外的解释力,同时还应考虑空间尺度及非生物因素的影响,为深入了解森林生产力的驱动机制提供科学依据.     

Temporal stability of aquatic food webs: partitioning the effects of species diversity, species composition and enrichment

Theory predicts that species diversity can enhance stability of community‐level biomass while simultaneously decreasing population‐level stability. Enrichment can theoretically destabilize communities but effects may become weaker with increasing diversity because of the inclusion of consumer‐resistant prey. Few experiments using direct manipulations of species diversity have tested these predictions. We used laboratory‐based aquatic food webs to examine the effects of species composition, diversity and enrichment on temporal variability of population‐ and community‐level biomass. We found weak effects of enrichment on population‐ and community‐level stability. However, diversity enhanced community‐level stability while species composition had no influence. In contrast, composition effects outweighed diversity effects when stability was measured at the population level. We found no negative effects of diversity on population‐level stability, in opposition to theory. Our results indicate that diversity can enhance stability in multitrophic systems, but effects vary with the scale of biological organization at which stability is measured.     

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.     

Enhanced interannual precipitation variability increases plant functional diversity that in turn ameliorates negative impact on productivity

Although precipitation interannual variability is projected to increase due to climate change, effects of changes in precipitation variance have received considerable less attention than effects of changes in the mean state of climate. Interannual precipitation variability effects on functional diversity and its consequences for ecosystem functioning are assessed here using a 6‐year rainfall manipulation experiment. Five precipitation treatments were switched annually resulting in increased levels of precipitation variability while maintaining average precipitation constant. Functional diversity showed a positive response to increased variability due to increased evenness. Dominant grasses decreased and rare plant functional types increased in abundance because grasses showed a hump‐shaped response to precipitation with a maximum around modal precipitation, whereas rare species peaked at high precipitation values. Increased functional diversity ameliorated negative effects of precipitation variability on primary production. Rare species buffered the effect of precipitation variability on the variability in total productivity because their variance decreases with increasing precipitation variance.     

Relationships between species diversity or community structure and productivity of woody-plants in a broad-leaved Korean pine forest in Jiaohe,Jilin, China

Aims Based on the dataset of a broad-leaved Korean pine forest in Jiaohe, Jilin Province, this research compared the influences of species diversity and community structure on productivity. We aim to explain the relationship between diversity and productivity for better forest management. Methods We used the data of 10 973 woody-plants in a 11.76 hm² large sample plot and analyzed the correlations between 7 different indices of species diversity or community structure and productivity. Structural equation model was used to compare the effects of species diversity and community structure on productivity. Important findings The results showed that: (1) Both species diversity and community structure had significant effects on productivity when they were considered separately in linear regression analysis, i.e. species evenness was negatively correlated with productivity, the Shannon index of community structure was positively correlated with productivity and the Gini index was negatively correlated with productivity. (2) In the structural equation model, when simultaneously considered, community structure had stronger influence on productivity than species diversity. Our research suggests that, the effects of community structure on productivity are greater than species diversity and it is important to increase community structure complexity to improve forest productivity during forest management.     

取样尺度对亚高寒草甸物种多样性与生产力关系的影响

植物物种多样性与生产力之间的关系是群落生态学的一个热点问题, 目前仍存在着很多争议。为探究自然群落中二者之间的关系, 对青藏高原亚高寒草甸3个样地的自然植物群落分别进行了不同取样面积的抽样调查。结果显示, 取样样地和取样尺度均对物种丰富度有显著影响, 取样样地而非取样尺度对群落地上生物量有显著性影响。在某一时刻对某一样地进行取样, 其单位面积生产力并不因取样面积的增加而提高, 而是保持恒定的, 尽管物种数随取样面积的增加而有明显增多。物种多样性与生产力之间的回归关系因样地与取样尺度不同而不同, 有U型、单峰型、正线性相关和无相关性, 其中无相关性出现的最多。据此推测, 亚高寒草甸群落物种多样性与生产力之间不存在某种确定性关系, 或者说, 亚高寒草甸物种多样性和生产力之间不存在必然的因果联系。     

刈割、施肥和浇水对高寒草甸物种多样性、功能多样性与初级生产力关系的影响

通过在高寒矮嵩草(Kobresia humilis)草甸为期4年的刈割(留茬1 cm、3 cm及不刈割)、施肥(尿素7.5 g·m^-2·a^-1+磷酸二铵1.8 g·m^-2·a^-1、不施肥)和浇水(20.1 kg·m^-2·a^-1、不浇水)控制实验, 对该生态系统中功能多样性与物种多样性的变化及其与初级生产力关系进行了研究。结果表明: 刈割和施肥显著影响植物性状和生产力, 而浇水作用微弱。刈割对物种多样性无影响, 但对不同功能多样性指数的影响不同; 施肥降低了物种多样性和以所有性状(除植株高度外)量化的功能多样性; 浇水仅对物种丰富度有微弱影响。各功能多样性的平均值能更好地反映群落物种间的功能差异。功能多样性和物种多样性呈正相关或不相关, 且二者与初级生产力的关系可表现为正相关、负相关和不相关, 说明三者间的关系是性状依赖的, 并受生境资源状况和刈割扰动的影响。植物功能性状对群落初级生产力有更加直接的影响。在考虑生物多样性对生态系统功能的影响时, 应更加注重对植物功能属性变化的研究。     

鄱阳湖湿地大型底栖无脊椎动物多样性对群落次级生产力及稳定性的影响

理解生物多样性对生态系统功能及稳定性的影响对于制定有效的保护管理策略有重要意义。然而,目前生物多样性与群落生产力、稳定性的关系仍存在争议。在鄱阳湖湿地布设30个采样点,于2019年秋季开展大型底栖无脊椎动物群落野外调查。基于底栖动物群落数据,采用广义加性模型分析物种、谱系、功能多样性对鄱阳湖湿地底栖动物群落次级生产力与稳定性的影响。结果表明:底栖动物群落的次级生产力与反映物种多样性的指数(Simpson多样性指数、Shannon-Wiener多样性指数、Pielou均匀度指数)、分类多样性指数、平均分类差异指数、功能丰富度指数等呈显著的负相关,其中Pielou均匀度指数与次级生产力的相关度最高(r²=0.33)。功能多样性对群落次级生产力的空间分异有最高的解释度(r²=0.75)。P/B值(次级生产力与生物量之比代表群落稳定性)与物种、谱系、功能多样性指数均呈正相关,其中功能丰富度与P/B值的相关度最高(r²=0.22)。反映物种多样性的三个指数总体上对P/B值的空间分异解释度最高(r²=0.37)。谱系多样性与次级生产力、P/B值的相关性相对较弱。生物多样性指数总体分别解释了次级生产力和群落稳定性中81.9%、54.8%的变异。上述研究结果表明,生物多样性与群落生产力、稳定性的关系可能因具体的生物群落而异。研究结果对于鄱阳湖湿地的底栖生物多样性保护有参考价值。     

吉林蛟河针阔混交林林层结构对生产力的影响

探讨不同林层物种多样性和林分结构与森林生产力之间的关系,为森林可持续经营管理策略优化提供理论支持。以吉林蛟河30 hm²针阔混交林固定监测样地中的38432株木本植物的野外调查数据为研究对象,利用结构方程模型探讨了不同林层物种多样性和林分结构多样性对森林生产力的作用机制。结果表明：（1）林冠层林分结构多样性和生产力之间呈负相关关系,路径系数为-0.138;林下层林分结构多样性和生产力之间呈正相关关系,路径系数为0.083。（2）林冠层物种多样性和生产力之间呈正相关关系,路径系数为0.099;林下层物种多样性和生产力之间关系不显著,且物种多样性在林冠层和林下层呈正相关关系,路径系数为0.147。物种多样性和林分结构多样性与森林生产力之间不存在唯一的关系,这种关系的大小、方向和潜在机制与林层有关。为此,在森林经营过程中要充分考虑不同林层物种多样性和林分结构多样性与森林生产力之间的关系。     

Functional and phylogenetic diversity determine woody productivity in a temperate forest

Understanding the relationships between biodiversity and ecosystem productivity has become a central issue in ecology and conservation biology studies, particularly when these relationships are connected with global climate change and species extinction. However, which facets of biodiversity (i.e. taxonomic, functional, and phylogenetic diversity) account most for variations in productivity are still not understood very well. This is especially true with regard to temperate forest ecosystems. In this study, we used a dataset from a stem‐mapped permanent forest plot in northeastern China exploring the relationships between biodiversity and productivity at different spatial scales (20 × 20 m; 40 × 40 m; and 60 × 60 m). The influence of specific environmental conditions (topographic conditions) and stand maturity (expressed by initial stand volume and biomass) were taken into account using the multivariate approach known as structural equation models. The variable “Biodiversity” includes taxonomic (Shannon), functional (FDis), and phylogenetic diversity (PD). Biodiversity–productivity relationships varied with the spatial scales. At the scale of 20 × 20 m, PD and FDis significantly affected forest biomass productivity, while Shannon had only indirect effects. At the 40 × 40 m and 60 × 60 m scales, biodiversity and productivity were weakly correlated. The initial stand volume and biomass were the most important drivers of forest productivity. The local environmental conditions significantly influenced the stand volume, biomass, biodiversity, and productivity. The results highlight the scale dependency of the relationships between forest biodiversity and productivity. The positive role of biodiversity in facilitating forest productivity was confirmed at the smaller scales. Our findings emphasize the fundamental role of environmental conditions in determining forest ecosystem performances. The results of this study provide a better understanding of the underlying ecological processes that influence specific forest biodiversity and productivity relationships.     

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。     

Climate change‐driven extinctions of tree species affect forest functioning more than random extinctions

Aim

Climate change affects forest functioning not only through direct physiological effects such as modifying photosynthesis and growing season lengths, but also through indirect effects on community composition related to species extinctions and colonizations. Such indirect effects remain poorly explored in comparison with the direct ones. Biodiversity–ecosystem functioning (BEF) studies commonly examine the effects of species loss by eliminating species randomly. However, species extinctions caused by climate change will depend on the species’ vulnerability to the new environmental conditions, thus occurring in a specific, non‐random order. Here, we evaluated whether successive tree species extinctions, according to their vulnerability to climate change, impact forest functions differently than random species losses.

Location

Eleven temperate forests across a gradient of climatic conditions in central Europe.

Methods

We simulated tree community dynamics with a forest succession model to study the impact of species loss on the communities’ aboveground biomass, productivity and temporal stability. Tree species were removed from the local pool (1) randomly, and according to (2) their inability to be recruited under a warmer climate or (3) their increased mortality under drier conditions.

Results

Results showed that non‐random species loss (i.e., based on their vulnerability to warmer or drier conditions) changed forest functioning at a different rate, and sometimes direction, than random species loss. Furthermore, directed extinctions, unlike random, triggered tipping points along the species loss process where forest functions were strongly impacted. These tipping points occurred after fewer extinctions in forests located in the coldest areas, where ecosystem functioning relies on fewer species.

Main conclusions

We showed that the extinction of species in a deterministic and mechanistically motivated order, in this case the species vulnerability to climate change, strengthens the selection effect of diversity on ecosystem functioning. BEF studies exploring the impact of species loss on ecosystem functioning using random extinctions thus possibly underestimate the potential effect of biodiversity loss when driven by a directional force, such as climate change.