Relationships between functional diversity and ecosystem functioning: A review

Functional diversity, which is the value, variation and distribution of traits in a community assembly, is an important component of biodiversity. Functional diversity is generally viewed as a key to understand ecosystem and community functioning. There are three components of functional diversity, i.e. functional richness, evenness and divergence. Functional diversity and species diversity can be either positively or negatively correlated, or uncorrelated, depending on the environmental conditions and disturbance intensity. Ecosystem functioning includes ecosystem processes, ecosystem properties and ecosystem stability. The diversity hypothesis and the mass ratio hypothesis are the two major hypotheses of explaining the effect of functional diversity on ecosystem functioning, diversity hypothesis reflects that organisms and their functional traits in a assemblage effect on ecosystem functioning by the complementarity of using resources, and mass ratio hypothesis emphasises the identify of the dominant species in a assemblage. These two hypotheses do not contradict each other and instead they reflect the two different sides of functional diversity and functional composition. The effect of functional diversity on ecosystem functioning also depends on abiotic factors, perturbation, management actions, etc. Function diversity potentially influences ecosystem service and management by effecting on ecosystem functioning. Ecosystem management groups should include functional diversity in their scheme and not just species richness.     

Predicting productivity in tropical reservoirs: The roles of phytoplankton taxonomic and functional diversity

Primary productivity is intimately linked with biodiversity and ecosystem functioning. Much of what is known today about such relationship has been based on the manipulation of species richness. Other facets of biodiversity, such as functional diversity, have been neglected within this framework, particularly in freshwater systems. We assess the adequacy of different diversity measures, from species richness and evenness, to functional groups richness and functional diversity indices, to predict primary productivity in 19 tropical reservoirs of central Brazil, built to generate hydroelectric energy. We applied linear mixed models (and model selection based on the Akaike’s information criterion) to achieve our goal, using chlorophyll-a concentration as a surrogate for primary productivity. A total of 412 species were collected in this study. Overall we found a positive relation between productivity and diversity, with functional evenness representing the only exception. The most parsimonious models never included functional group classifications, with at least one continuous measure of functional diversity being present in many models. The best model included only species richness and explained 24.1% of variability in productivity. We therefore advise the use of species richness as an indicator of productivity in tropical freshwater environments. However, since the productivity–diversity relationship is known to be scale dependent, we recommend the use of continuous measures of functional diversity in future biodiversity and ecosystem functioning studies, in order to be certain that all functional differences between communities are being accounted for.     

Effects of different components of diversity on productivity in artificial plant communities

In an experiment on artificial plant communities, the effects of three components of plant diversity—plant species diversity, plant functional group diversity and plant functional diversity—on community productivity and soil water content were compared. We found that simple regression analysis showed a positive diversity effect on ecosystem processes (productivity and soil water content). However, when three components of diversity were included in the multiple regression analyses, the results showed that functional group diversity and functional diversity had more important effects on productivity and resource use efficiency. These results suggested that, compared with species number, functional differences among species and the range of functional traits carried by plants are the basis of biodiversity effects on ecosystem functioning. These diversity effects of increasing functional group diversity or functional diversity were likely because species differing greatly in size, life form, phenology and capacity to capture and use resources efficiently in diverse communities realize complementary resource use in temporal, spatial, and biological ways.     

种、种的多样性及退化生态系统功能的恢复和维持研究

物种多样性是生态系统的重要特征并维持系统的功能支行,生物种和不同种类构成的群落为人类提供诸如营养物质循环、生物生产力、营养功能等形式的重要生态服务,特种多样性与生态系统抵御逆境和干扰的能力紧密相关,多样性的提高会增加系统的稳定性,与单个种和种类的数量相比,功能群和功能多样性对生态系统功能的影响效应要大得多,且易于被用来测度稳定性和预测群落变化,本文提出并探讨了种对生态系统功能作用的几种形式,理解物种多样性与生态系统的功能关系能指导退化生态系统恢复和维持其功能的实践活动,尤其为恢复的初始阶段进行群落的“种类组装”提供生态理论基础。     

Partitioning taxonomic diversity of aquatic insect assemblages and functional feeding groups in neotropical savanna headwater streams

Biological diversity can be divided into: alpha (α, local), beta (β, difference in assemblage composition among locals), and gamma (γ, total diversity). We assessed the partitioning of taxonomic diversity of Ephemeroptera, Plecoptera and Trichoptera (EPT) and of functional feeding groups (FFG) in neotropical savanna (southeastern Brazilian cerrado) streams. To do so, we considered three diversity components: stream site (α), among stream sites (β₁), and among hydrologic units (β₂). We also evaluated the association of EPT genera composition with heterogeneity in land use, instream physical habitat structure, and instream water quality variables. The percentage of EPT taxonomic α diversity (20.7%) was smaller than the β₁ and β₂ diversity percentages (53.1% and 26.2%, respectively). The percentage of EPT FFG collector-gatherer α diversity (26.5%) was smaller than that of β₁ diversity (55.8%) and higher than the β₂ (17.7%) diversity. The collector-gatherer FFG was predominant and had the greatest β diversity percentage among stream sites (β₁, 55.8%). Our findings support the need for implementing regional scale conservation strategies in the cerrado biome, which has been degraded by anthropogenic activities.     

松嫩平原南部主要群落植物多样性的比较研究

应用Ｍａｒｇａｌｅｆ（Ｒ１）和Ｍｅｎｈｉｎｉｃｋ（Ｒ２）丰富度指数、Ｈｉｌｌ多样性指数（Ｎ１,Ｎ２）和Ｐｉｅｌｏｕ均匀度指数（Ｅ）比较研究了松嫩平原南部１４个主要群落的植物多样性、生活型多样性及植物多样性与群落演替的关系结果表明,分布于低平原的羊草－蔓萎陵菜群落的植物多样性和丰富度指数最大,Ｒ１为１１．９６,Ｒ２为５．２４,Ｎ１为１３．９２,Ｎ２为８．７６,分布于固定砂丘的榆树疏林和贝加尔针茅等４个群落有较高的植物多样性,而分布于碱化草甸的角碱蓬等群落植物多样性最小,Ｒ１为１．３１,Ｒ２为０７１,Ｎ１为１．８４,Ｎ２为１．４６．具有较高植物多样性的群落其植物生活型多样性指数也较高．在演替系列中,羊草群落的植物多样性指数介于演替初期和中期阶段之间．     

祁连山区不同海拔草地群落的物种多样性

依据植物在各海拔带出现的频率和不同海拔带植物种的生活型,对祁连山区草地群落海拔2500～3000m的物种α多样性变化特征进行了研究．结果表明,随海拔升高,Shahnon-Wiener指数从1．432上升到1．832,Simpson指数的变化较为复杂．海拔2701～2800m带是物种多样性的一个重要转折点,是物种多样性较丰富的地带．在祁连山草地群落中,有毒的与食性较差的物种在各海拔带所占比例均超过50％,生物量占到整个样地生物量的60％以上,物种多样性已呈现衰退．     

新疆博格达山地面生苔藓植物物种多样性研究

采用相似性系数和物种多样性指数对新疆博格达山地面生苔藓植物的物种多样性进行了研究．结果表明,博格达山地区苔藓植物区系具有丰富性和复杂性的特点,共有地面生苔藓植物32科73属186种(含变种、变型);山地荒漠带和山地草原带苔藓植物物种相似性最高,为0．6809,山地森林带和高山垫状植被带相似性最低,仅为0．1342;山地森林带苔藓植物在整个博格达山地面生苔藓植物群落中占据优势地位;山地森林带苔藓植物物种多样性最为丰富,为该山地苔藓植物多样性的分布中心,是苔藓植物多样性保护的关键地区．     

Exploring the functional significance of forest diversity: A new long-term experiment with temperate tree species (BIOTREE)

Effects of biodiversity on ecosystem functioning have been mainly studied in experiments that artificially create gradients in grassland plant diversity. Woody species were largely excluded from these early experiments, despite the ecological and socioeconomic importance of forest ecosystems. We discuss conceptual aspects of mechanistically driven research on the biodiversity–ecosystem functioning relationship in forests, including the comparison of scientific approaches like ‘observational studies’, ‘removal experiments’, and ‘synthetic-assemblage experiments’. We give a short overview on the differences between herbaceous and forest ecosystems, focusing on canopy characteristics, and the possibilities for individual versus population-based investigations.We present detailed information about the first large-scale, multisite and long-term biodiversity–ecosystem functioning experiment with tree species of temperate forests (BIOTREE – BIOdiversity and ecosystem processes in experimental TREE stands). At three sites of differing geology and local climate, we planted 200,000 saplings on a total area of 70 ha. At two sites, diversity gradients were established by varying the number of tree species (BIOTREE-SPECIES). At a third site, only functional diversity at a constant level of tree species richness was manipulated by selecting mixtures that differ in the functional trait values of the corresponding species (BIOTREE-FD). Additional experimental treatments at the subplot level include silvicultural management options, the addition of subdominant species, and the reduction of genetic diversity. Response variables focus on productivity, biogeochemical cycles and carbon sequestration, and resource use complementarity.We explore the use of different measures of functional diversity for a posteriori classifications of functional richness and their use in the analysis of our tree diversity experiment. The experiment is thought to provide a long-term research platform for a variety of scientific questions related to forest biodiversity and ecosystem processes.     

Effects of plant diversity on invertebrate herbivory in experimental grassland

The rate at which a plant species is attacked by invertebrate herbivores has been hypothesized to depend on plant species richness, yet empirical evidence is scarce. Current theory predicts higher herbivore damage in monocultures than in species-rich mixtures. We quantified herbivore damage by insects and molluscs to plants in experimental plots established in 2002 from a species pool of 60 species of Central European Arrhenatherum grasslands. Plots differed in plant species richness (1, 2, 4, 8, 16, 60 species), number of functional groups (1, 2, 3, 4), functional group and species composition. We estimated herbivore damage by insects and molluscs at the level of transplanted plant individuals (“phytometer” species Plantago lanceolata, Trifolium pratense, Rumex acetosa) and of the entire plant community during 2003 and 2004. In contrast to previous studies, our design allows specific predictions about the relative contributions of functional diversity, plant functional identity, and species richness in relation to herbivory. Additionally, the phytometer approach is new to biodiversity-herbivory studies, allowing estimates of species-specific herbivory rates within the larger biodiversity-ecosystem functioning context. Herbivory in phytometers and experimental communities tended to increase with plant species richness and the number of plant functional groups, but the effects were rarely significant. Herbivory in phytometers was in some cases positively correlated with community biomass or leaf area index. The most important factor influencing invertebrate herbivory was the presence of particular plant functional groups. Legume (grass) presence strongly increased (decreased) herbivory at the community level. The opposite pattern was found for herbivory in T. pratense phytometers. We conclude that (1) plant species richness is much less important than previously thought and (2) plant functional identity is a much better predictor of invertebrate herbivory in temperate grassland ecosystems.     

Land-use intensification reduces functional redundancy and response diversity in plant communities

Ecosystem resilience depends on functional redundancy (the number of species contributing similarly to an ecosystem function) and response diversity (how functionally similar species respond differently to disturbance). Here, we explore how land-use change impacts these attributes in plant communities, using data from 18 land-use intensity gradients that represent five biomes and > 2800 species. We identify functional groups using multivariate analysis of plant traits which influence ecosystem processes. Functional redundancy is calculated as the species richness within each group, and response diversity as the multivariate within-group dispersion in response trait space, using traits that influence responses to disturbances. Meta-analysis across all datasets showed that land-use intensification significantly reduced both functional redundancy and response diversity, although specific relationships varied considerably among the different land-use gradients. These results indicate that intensified management of ecosystems for resource extraction can increase their vulnerability to future disturbances.
Ecology Letters (2010) 13: 76–86     

Cascading extinctions, functional complementarity, and selection in two-trophic-level model communities: A trait-based mechanistic approach

The influence of diversity on ecosystem functioning and ecosystem services is now well established. Yet predictive mechanistic models that link species traits and community-level processes remain scarce, particularly for multitrophic systems. Here we revisit MacArthur's classical consumer resource model and develop a trait-based approach to predict the effects of consumer diversity on cascading extinctions and aggregated ecosystem processes in a two-trophic-level system. We show that functionally redundant efficient consumers generate top-down cascading extinctions. This counterintuitive result reveals the limits of the functional redundancy concept to predict the consequences of species deletion. Our model also predicts that the biodiversity-ecosystem functioning relationship is different for different ecosystem processes and depends on the range of variation of consumer traits in the regional species pool, which determines the sign of selection effects. Lastly, competition among resources and consumer generalism both weaken complementarity effects, which suggests that selection effects may prevail at higher trophic levels. Our work emphasizes the potential of trait-based approaches for transforming biodiversity and ecosystem functioning research into a more predictive science.     

历山森林群落物种多样性与群落结构研究

采用多样性指数、丰富度指数和均匀度指数对山西历山森林群落物种多样性进行了研究.结果表明,各群落类型的物种多样性指数的顺序为：青榨槭＋五角枫林>青榨槭＋鹅耳枥林>辽东栎林>栓皮栎林>华山松＋红桦林>油松＋栎林>白桦林>侧柏＋栓皮栎林>红桦林>华山松林>侧柏林>油松林>红桦＋山杨林;森林群落灌木层多样性指数和丰富度指数大于草本层和乔木层,而均匀度指数在乔木层、灌木层和草本层则表现出多样化的趋势;海拔1 000～1 920 m之间,Shannon-Wiener指数、Hill多样性指数及物种丰富度与海拔梯度均呈正相关(P<0.05),海拔大于1 920 m则呈现负相关;这些指数沿海拔梯度的变化呈现出“中间高度膨胀”的规律,即中等海拔高度上物种多样性高而低海拔和高海拔物种多样性低.这主要是由于在中海拔（约1 900 m）地段水热条件组合较好,人类活动干扰较少所致.     

Overyielding and the role of complementary use of nitrogen in wetland plant communities

Biodiversity and ecosystem functioning experiments have demonstrated that plant biomass of species grown in mixtures is often greater than plant biomass of monocultures (i.e., mixtures over yield). While we understand that plant species utilize resources differently, how a combination of species increases resource use and productivity is not well known, especially in wetland ecosystems. Here, we used a mesocosm experiment to explore diversity effects on plant biomass production and to examine the role of N partitioning as a mechanism for overyielding in wetland ecosystems. Plant functional groups (FGs) represented the unit of diversity, and we included five levels of diversity (0-4 FGs). To test for N partitioning, we used a stable isotope technique to determine niche breadth and proportion similarity of inorganic N use (NO₃⁻ and NH₄⁺) for individual FGs as well as mixtures containing 3 and 4 FGs. We found that total plant biomass increased in the first season from an average of 290 ± 60 SE g ash-free dry mass (AFDM) m⁻² at the 1 FG level to 490 ± 70 g AFDM m⁻² at the 4 FG level and in the second season from an average of 560 ± 80 g AFDM m⁻² at the 1 FG level to 1000 ± 90 g AFDM m⁻² at the 4 FG level indicating overyielding. Plant species comprising the majority of mesocosm biomass demonstrated preferential uptake of ¹⁵NO₃⁻, while species with relatively less biomass (e.g., Acorus calamus and Carex crinita) preferred ¹⁵NH₄⁺. Concentrations of ¹⁵N in biomass increased with FG richness, but only in the ¹⁵NO₃⁻ treatment. Niche breadth did not vary among levels of FG richness. We observed a greater niche overlap with an increase of FGs, with species taking up greater proportion of ¹⁵NO₃⁻ than ¹⁵NH₄⁺. Our results indicate that plant overyielding in wetland mesocosms is not the result of niche partitioning of N chemical forms, but is associated with greater uptake of NO₃.     

The role of phytoplankton diversity metrics in shallow lake and river quality assessment

Ecological water quality problems are frequently connected to increment of phytoplankton productivity and overdominance of some phytoplankton species. Metrics that show monotonously increasing or decreasing tendencies along stressor gradients is recommended for ecological state assessment. Diversity metrics are influenced by various physical disturbances and show high within-year variability; thus, there is no agreement on the usefulness of these metrics as state indicators.To test the usefulness of phytoplankton diversity in ecological state assessment we investigated the productivity–diversity relationships for lakes and rivers in the Carpathian Basin (Hungary). We demonstrated that the shape of productivity–diversity relationship depends on the investigated water body type. Regarding lakes, hump-shaped relationship was found for all computed metrics. Parallel with the increase in phytoplankton productivity values, diversity metrics showed monotonously increasing tendencies in rhithral and decreasing tendencies in large potamal rivers. We found no systematic relationship in the case of small lowland rivers.Changes of diversity metrics calculated for species and functional groups showed similar tendencies within the types, only the slopes of regression lines differ each other.The use of diversity metrics as ecological state indicators should be restricted to water body types where diversity decreases or increases monotonously with phytoplankton biomass. Regarding the lakes the use of diversity metrics is not recommended for ecological state assessment. In rhithral and large potamal river assessment, application of diversity metrics should be strongly considered. We demonstrated that diversity metrics can be useful components of multimetric indices proposed to use by the Water Framework Directive.     

外来植物凋落物分解对底栖动物多样性及其摄食功能群的影响

在亚热带地区一条2级溪流中比较了本地植物(红锥和薏米)和外来植物(托里桉和马缨丹)凋落物的分解速率及底栖动物定殖及其摄食功能群的组成。结果显示,本地植物凋落物的分解速率为:0.020 d-1(红锥)和0.056 d-1(薏米),外来树叶的分解速率则为:0.038 d-1(托里桉)和0.041 d-1(马缨丹),它们均属于快速分解组。定殖在外来和本地树叶凋落物上的大型底栖动物种类分别为21种和24种。在外来和本地树叶上定殖的底栖动物平均密度差异不显著(P0.05)。从大型底栖动物摄食功能群的比例来看,定殖在本地凋落物上的集食者的比例最高(73.5%),其次是撕食者(10.9%)和捕食者(8.7%),刮食者(6.8%)的比例最低,在外来凋落物上,摄食功能群的比例分别为集食者(67.6%)、撕食者(13.9%)、刮食者(12.1%)和捕食者(6.1%),两者相比,底栖动物的摄食功能群均以集食者和撕食者为主,在组成比例上并无显著差异。结果表明2种外来植物凋落物分解对底栖动物多样性及其摄食功能群的影响不明显。     

Functional diversity research of forest communities in the Xiaowutai Mountain National Nature Reserve,Hebei

Plant functional traits are the plant physiological characteristics which can response to the changes of the living environment and have a certain impact on the ecosystem structure and function. The objective of our study was to explore characters of present functional diversity indices, the relationships between functional diversity and environmental variables, the relevance of species diversity and functional diversity. In this paper, habitat type, seed dispersal, pollination method, life cycle, life form, leaf form, leaf hair type, flowering period and flowering time were chosen as functional traits, and the research were done in the typical forest communities in the Xiaowutai Mountain National Nature Reserve, Hebei. One hundred and forty-eight quadrats (10 m × 10 m) of forest communities were established along altitude gradients, at the same time, species composition, functional traits, and environmental variables were measured in each quadrat. The results showed that functional diversity indices in forest communities that were calculated by functional distances varied greatly. Functional diversity indices (FAD, MFAD, FDp, FDc, FRic, Rao and FDis) had highly significantly positive correlation with Patrick index and showed a linear increasing trend. All the nine functional diversity indices (FAD, MFAD, FDp, FDc, FRic, Rao, FEve, FDiv, FDis) had significantly correlation with Shannon-Wiener index and Pielou index. Only FDiv showed significantly negative correlation, and the other eight functional diversity indices showed positive correlation. Environmental filtering was important to functional diversity pattern, and functional diversity indices showed correlation with environmental variables. Altitude was a significant factor to functional diversity in forest communities. Except for FDiv, other functional diversity indices displayed a decreasing trend along altitude gradients. Among all the functional diversity indices, only Rao and FDis showed significantly positive correlation with aspect. The functional diversity indices (FAD, MFAD, FDp, FDc, FRic, Rao and FDdis) showed a negative correlation with slope, slope position, litter layer thickness, soil thickness, while, they showed a positive correlation with soil temperature and disturbance. All the nine indices were proved successful in the analysis of functional diversity in forest communities with different effectiveness. They were divided into three categories, functional richness (FAD, MFAD, FDp, FDc, FRic), functional divergence (Rao, FDis), functional evenness (FEve, FDis). Meanwhile, each category was highly inter-correlated and each category was relatively independent with other categories. The study of functional diversity provides a number of ecological indication and monitoring methods for the forest, and it can address a wide range of important ecological questions that links species and ecosystems through mechanisms in biodiversity research.     

Biodiversity and resilience of arthropod communities after fire disturbance in temperate forests

Changes in ecosystem functions following disturbances are of central concern in ecology and a challenge for ecologists is to understand the factors that affect the resilience of community structures and ecosystem functions. In many forest ecosystems, one such important natural disturbance is fire. The aim of this study was to understand the variation of resilience in six functional groups of invertebrates in response to different fire frequencies in southern Switzerland. We measured resilience by analysing arthropod species composition, abundance and diversity in plots where the elapsed time after single or repeated fires, as determined by dendrochronology, varied. We compared data from these plots with data from plots that had not burned recently and defined high resilience as the rapid recovery of the species composition to that prior to fire. Pooling all functional groups showed that they were more resilient to single fires than to repeated events, recovering 6–14 years after a single fire, but only 17–24 years after the last of several fires. Flying zoophagous and phytophagous arthropods were the most resilient groups. Pollinophagous and epigaeic zoophagous species showed intermediate resilience, while ground-litter saprophagous and saproxylophagous arthropods clearly displayed the lowest resilience to fire. Their species composition 17–24 years post-burn still differed markedly from that of the unburned control plots. Depending on the fire history of a forest plot, we found significant differences in the dominance hierarchy among invertebrate species. Any attempt to imitate natural disturbances, such as fire, through forest management must take into account the recovery times of biodiversity, including functional group composition, to ensure the conservation of multiple taxa and ecosystem functions in a sustainable manner.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .