Conservation implications of the link between biodiversity and ecosystem functioning

The relationship between biodiversity and individual ecosystem processes is often asymptotic, saturating at relatively low levels, with some species contributing more strongly than others. This has cast doubt on arguments for conservation based on maintenance of the functioning of ecosystems. However, we argue that the link between biodiversity and ecosystem functioning is an important additional argument for conservation for several reasons. (1) Although species differ in importance to ecosystem processes, we do not believe that this argues for preservation of just a few species for two reasons: first, it is nearly impossible to identify all species important to the numerous systems and processes on which humans depend; second, the important species themselves may depend on an unknown number of other species in their communities. (2) Arguments for conservation based on ecosystem functioning are complementary to other utilitarian, ethical and aesthetic justifications. No single reason will convince all people or protect all species, however the combination produces a strong case for conservation of biodiversity. (3) Even if the relationship between biodiversity and ecosystem functioning is asymptotic at local spatial scales and in the short term, effects of biodiversity loss are likely to be important at larger temporal and spatial scales. (4) Initial arguments for the importance of biodiversity for ecosystem functioning were largely based on a precautionary approach (points 1-3). However, we are now moving to a scientific position based on accumulating experimental evidence. The future challenge is the integration of this scientific research with policy.     

The effects of thin mud deposits on the behaviour of a deposit-feeding tellinid bivalve: implications for ecosystem functioning

A laboratory experiment was conducted, to examine how feeding, siphon activity and movement behaviours of a tellinid bivalve, Macomona liliana (Iredale, 1915), were affected by thin surface layers of mud (1–4 mm) and the incorporation of mud into surface (0–2 cm) sediment. Time-lapse photography and porewater pressure sensors were used to characterise changes in behaviour before and after mud addition. Mud addition modified the sediment matrix which had an immediate effect on M. liliana behaviour; significantly affecting the rates of feeding and the degree of porewater pressurisation during feeding events. Surface activity indicated maintenance and clearance of established burrow structures and increased ventilation. Ultimately, subtle modifications in behaviour and sediment hydraulic conductivity may have consequences for nutrient exchange and benthic-pelagic coupling. Our results emphasise the need to consider the impacts of low level stressors when they affect the behaviour of structurally important species.     

Temporal niches of shredders in lake littorals with possible implications on ecosystem functioning

Temporal growth separation of shredders is known in streams but has not been reported from lakes. In the present study, temporal niche/trait differentiations among shredders in lakes were investigated. We sampled quantitatively three lakes in SE Sweden over a period of 18 months. Dry weight and number of individuals of the collected shredders were measured monthly. Standing stock of detritus types was also monitored. The same 10 species of lentic shredders were found in each lake, one isopod and larvae of nine trichopterans. Functionally, the shredders could be categorized into two main groups; winter and summer growing species. However, also within these groups, temporal differences in growth pattern existed. The main input of detritus occurred during the autumnal leaf fall and a majority of winter shredders had the start of their lives tied to this period. A succession in loss of detritus types was evident with easily degraded matter disappearing first followed by more resistant matter. Shredder species richness, shredder biomass per m² and the ratio coarse/fine detritus all reached its annual low in late summer. We propose a temporal link between the shredder groups and the organic matter subject to decomposition; the successive palatability of coarse detritus is likely to make a temporally separated community of shredders efficient in terms of decomposition. We believe that a temporal differentiation per se is sufficient to conclude that different impact on ecosystem function exists among shredders. Additionally we discuss impacts of differences in abundance and shredding capacity among the species.     

Biodiversity and ecosystem functioning decoupled: invariant ecosystem functioning despite non‐random reductions in consumer diversity

Most research that demonstrates enhancement and stabilization of ecosystem functioning due to biodiversity is based on biodiversity manipulations within one trophic level and measuring changes in ecosystem functions provided by that same trophic level. However, it is less understood whether and how modifications of biodiversity at one trophic level propagate vertically to affect those functions supplied by connected trophic levels or by the whole ecosystem. Moreover, most experimental designs in biodiversity–ecosystem functioning research assume random species loss, which may be of little relevance to non‐randomly assembled communities. Here, we used data from a published ecotoxicological experiment in which an insecticide gradient was applied as an environmental filter to shape consumer biodiversity. We tested how non‐random consumer diversity loss affected gross primary production (an ecosystem function provided by producers) and respiration (an ecosystem function provided by the ecosystem as whole) in species‐rich multitrophic freshwater communities (total of 128 macroinvertebrate and 59 zooplankton species across treatments). The insecticide decreased and destabilized macroinvertebrate and, to a lesser extent, zooplankton diversity. However, these effects on biodiversity neither affected nor destabilized any of the two studied ecosystem functions. The main reason for this result was that species susceptible to environmental filtering were different from those most strongly contributing to ecosystem functioning. The insecticide negatively affected the most abundant species, whereas much less abundant species had the strongest effects on ecosystem functioning. The latter finding may be explained by differences in body size and feeding guild membership. Our results indicate that biodiversity modifications within one trophic level induced by non‐random species loss do not necessarily translate into changes in ecosystem functioning supported by other trophic levels or by the whole community in the case of limited overlap between sensitivity and functionality.     

Biodiversity and ecosystem functioning: It is time for dispersal experiments

The experimental study of the relationship between biodiversity and ecosystem function has mainly addressed the effect of species and number of functional groups. In theory, this approach has mainly focused on how extinction affects function, whereas dispersal limitation of ecosystem function has been rarely discussed. A handful of seed introduction experiments, as well as numerous observations of the effects of long‐distance dispersal of alien species, indicate that ecosystem function may be strongly determined by dispersal limitation at the local, regional and/or global scales. We suggest that it is time to replace biodiversity manipulation experiments, based on random draw of species, with those addressing realistic scenarios of either extinction or dispersal. Experiments disentangling the dispersal limitation of ecosystem function should have to take into account the probability of arrival. The latter is defined as the probability that a propagule of a particular species will arrive at a particular community. Arrival probability depends on the dispersal ability and the number of propagules of a species, the distance a species needs to travel, and the permeability of the matrix landscape. Current databases, in particular those in northwestern and central Europe now enable robust estimation of arrival probability in plant communities. We suggest a general hypothesis claiming that dispersal limitation according to arrival probability will have ecosystem‐level effects different from those arising due to random arrival. This hypothesis may be rendered more region‐, landscape‐ or ecosystem‐specific by estimating arrival probabilities for different background conditions.     

Biodiversity and ecosystem functioning: recent theoretical advances

The relationship between biodiversity and ecosystem functioning has emerged as a major scientific issue today. As experiments progress, there is a growing need for adequate theories and models to provide robust interpretations and generalisations of experimental results, and to formulate new hypotheses. This paper provides an overview of recent theoretical advances that have been made on the two major questions in this area: (1) How does biodiversity affect the magnitude of ecosystem processes (short‐term effects of biodiversity)? (2) How does biodiversity contribute to the stability and maintenance of ecosystem processes in the face of perturbations (long‐term effects of biodiversity)?
Positive short‐term effects of species diversity on ecosystem processes, such as primary productivity and nutrient retention, have been explained by two major types of mechanisms: (1) functional niche complementarity (the complementarity effect), and (2) selection of extreme trait values (the selection effect). In both cases, biodiversity provides a range of phenotypic trait variation. In the complementarity effect, trait variation then forms the basis for a permanent association of species that enhances collective performance. In the selection effect, trait variation comes into play only as an initial condition, and a selective process then promotes dominance by species with extreme trait values. Major differences between within‐site effects of biodiversity and across‐site productivity–diversity patterns have also been clarified. The local effects of diversity on ecosystem processes are expected to be masked by the effects of varying environmental parameters in across‐site comparisons.
A major reappraisal of the paradigm that has dominated during the last decades seems necessary if we are to account for long‐term effects of biodiversity on ecosystem functioning. The classical deterministic, equilibrium approaches to stability do not explain the reduced temporal variability of aggregate ecosystem properties that has been observed in more diverse systems. On the other hand, stochastic, nonequilibrium approaches do show two types of biodiversity effects on ecosystem productivity in a fluctuating environment: (1) a buffering effect, i.e., a reduction in the temporal variance; and (2) a performance‐enhancing effect, i.e., an increase in the temporal mean. The basic mechanisms involved in these long‐term insurance effects are very similar to those that operate in short‐term biodiversity effects: temporal niche complementarity, and selection of extreme trait values. The ability of species diversity to provide an insurance against environmental fluctuations and a reservoir of variation allowing adaptation to changing conditions may be critical in a long‐term perspective.
These recent theoretical developments in the area of biodiversity and ecosystem functioning suggest that linking community and ecosystem ecology is a fruitful avenue, which paves the way for a new ecological synthesis.     

Effects of vegetation state on biodiversity and nitrogen retention in created wetlands: a test of the biodiversity–ecosystem functioning hypothesis

1. Nitrogen retention in wetlands provides an example of an ecosystem function that is desired by human society, and is a rationale for the creation of wetlands to decrease nitrogen fluxes from nitrate‐loaded river catchments to coastal waters. 2. Here, we tested the impact of different vegetation states on species diversity and nitrogen retention during 4 years in surface‐flow wetlands receiving nitrate‐rich water. Tall emergent vegetation or submerged vegetation was introduced to six experimental wetlands each and six wetlands were left as unplanted controls for free development of vegetation. This resulted in three vegetation states dominated by emergent vegetation, by a mixture of submerged vegetation and filamentous green algae or by filamentous green algae. 3. Species diversity (species richness and Shannon diversity) of plants was initially lowest in free development wetlands, but during the study became lower in the emergent vegetation wetlands than in the other wetlands. Diversity of macroinvertebrates was initially lower in the submerged vegetation wetlands than in the other wetlands, but this difference disappeared during the study. Nitrogen retention was consistently higher in emergent vegetation wetlands than in the other wetlands throughout the study. 4. We conclude that plant diversity in wetlands dominated by tall emergent vegetation gradually became lower than in other wetlands, due to dominant species competitively excluding other plants. However, these wetlands were more efficient at removing nitrogen than those dominated by filamentous algae or submerged macrophytes. 5. Management of wetlands often aims to decrease the dominance of tall emergent vegetation for the benefit of plant species diversity and habitat heterogeneity. Our results demonstrate a biodiversity benefit, but also show that this strategy may decrease the ability of wetlands to remove nitrogen. In this case, there is no support for the hypothesis that biodiversity enhances ecosystem function.     

Manganese accumulation in rice: implications for photosynthetic functioning

In order to gain fundamental insights into the nature of the adaptation to Mn excess, the characterisation of the photosynthetic apparatus in Mn-treated rice was carried out in 21-day-old plants. We found 17- and 11-fold increases in Mn in the leaf tissues and in thylakoid, respectively, when the plants were grown hydroponically in nutrient solutions with Mn concentrations between 0.125 and 32 mg l(-1) (2.3 and 582.5 microM). Net photosynthesis and the photosynthetic capacity decreased after the 0.5 and 2 mg l(-1) (9.1 and 36.4 microM) Mn treatment, respectively. The stomatal conductance displayed a similar trend to that of photosynthetic capacity. The levels of basal chlorophyll fluorescence and the ratio between variable and maximum chlorophyll fluorescence did not vary significantly among treatments, but the photochemical quenching and the quantum yield of non-cyclic electron transport increased until the 2 mg l(-1) (36.4 microM) Mn treatment. The lipid matrix of thylakoids revealed a global increase in the proportions of phospholipids, relative to galactolipids. This pattern was coupled with diminishing levels of monogalactosyldiacylglycerol. The relative ratio between total carotenoids and total chlorophylls decreased until the last Mn treatment, yet the levels of carotenes, zeaxanthin, and violaxanthin plus antheraxanthin displayed different patterns. It was further found that the de-epoxidation state involving the components of the xanthophylls cycle increased until the 8 mg l(-1) (145.6 microM) Mn treatment. The levels of the photosynthetic electron carriers displayed different patterns, with plastocyanin and the high and low forms of cytochrome b559 remaining steady, whereas cytochromes b563 and f increased until the 8 mg l(-1) (145.6 microM) Mn treatment and the quinone pool increased until the highest Mn treatment. It was concluded that Mn-mediated inhibition of rice photosynthesis barely implicates stomatal conductance, as well as the distribution of energy within the photosystems. In this context, alterations to the relative proportions of the different acyl lipids and isoprenoids, as well as to the accumulations of the photosynthetic electron carriers, seem to play a major role.     

Warming reduces the growth and diversity of biological soil crusts in a semi-arid environment: implications for ecosystem structure and functioning

Biological soil crusts (BSCs) are key biotic components of dryland ecosystems worldwide that control many functional processes, including carbon and nitrogen cycling, soil stabilization and infiltration. Regardless of their ecological importance and prevalence in drylands, very few studies have explicitly evaluated how climate change will affect the structure and composition of BSCs, and the functioning of their constituents. Using a manipulative experiment conducted over 3 years in a semi-arid site from central Spain, we evaluated how the composition, structure and performance of lichen-dominated BSCs respond to a 2.4°C increase in temperature, and to an approximately 30 per cent reduction of total annual rainfall. In areas with well-developed BSCs, warming promoted a significant decrease in the richness and diversity of the whole BSC community. This was accompanied by important compositional changes, as the cover of lichens suffered a substantial decrease with warming (from 70 to 40% on average), while that of mosses increased slightly (from 0.3 to 7% on average). The physiological performance of the BSC community, evaluated using chlorophyll fluorescence, increased with warming during the first year of the experiment, but did not respond to rainfall reduction. Our results indicate that ongoing climate change will strongly affect the diversity and composition of BSC communities, as well as their recovery after disturbances. The expected changes in richness and composition under warming could reduce or even reverse the positive effects of BSCs on important soil processes. Thus, these changes are likely to promote an overall reduction in ecosystem processes that sustain and control nutrient cycling, soil stabilization and water dynamics.     

The significance of tree-tree interactions for forest ecosystem functioning

Global change exposes forest ecosystems to many risks including novel climatic conditions, increased frequency of climatic extremes and sudden emergence and spread of pests and pathogens. At the same time, forest landscape restoration has regained global attention as an integral strategy for climate change mitigation. Owing to unpredictable future risks and the need for new forests that provide multiple ecosystem services, mixed-species forests have been advocated for this purpose. However, the successful establishment of mixed forests requires intrinsic knowledge of biodiversity's role for forest ecosystem functioning. In this respect, a better understanding of tree-tree interactions and how they contribute to observed positive tree species richness effects on key ecosystem functions is critical. Here, we review the current knowledge of the underlying mechanisms of tree-tree interactions and argue that positive net biodiversity effects at the community scale may emerge from the dominance of positive over negative interactions at the local neighbourhood scale. In a second step, we demonstrate how tree-tree interactions and the immediate tree neighbourhood's role can be systematically assessed in a tree diversity experiment. The expected results will improve predictions about the effects of tree interactions on ecosystem functioning based on general principles. We argue that this knowledge is urgently required to guide the design of tree species mixtures for the successful establishment of newly planted forests.     

生物多样性与生态系统功能:进展与争论

生物多样性与生态系统功能的关系已成为当前人类社会面临的一个重大科学问题,生物多样性的空前丧失,促使人们开展了大量研究工作来描述物种多样性-生态系统功能关系,并试图揭示多样性与系统功能关系的内在机制,本文将多样性对生态系统功能作用机制的有关假说分为统计学与生物学两大类：前者是从统计学角度来解释观察到的多样性-系统功能模式,包括抽样效应,统计均衡效应等;而后者是基于多样性的生物学效应给出的,包括生态位互补,种间正相互作用,保险效应等,本文较为详细地介绍了该领域内有代表性的实验工作,包括“生态箱”实验,Cedar Creek草地多样性实验,微宇宙实验,欧洲草地实验,以及在这些实验结果解释上的激烈争论。     

With the worms: Soil biodiversity and ecosystem functioning

The role of soil as a carbon sink, nutrient recycler and pollutant remover is becoming increasingly apparent. But it is the animals and microbes that live in the soil that enable all these activities. Do we know enough about these systems to prevent permanent damage?     

生物多样性与生态系统功能:最新的进展与动向

生物多样性与生态系统功能的关系及其内在机制是当前生态学领域的重大科学问题。 2 0 0 2年以来人们不再过多地纠缠于“抽样 -互补之争” ,对这一世纪课题的认识又有了新的进展。 (1)人们开始运用已有的知识揭示更大时间和空间尺度上的物种多样性 -生态系统功能关系。多样性作用机制可能存在着动态变化———“抽样向互补转型” :群落建立初期 ,抽样效应是主要的多样性作用机制 ;随时间推移 ,生态位互补成为主要机制。理论研究则预测 :局域尺度上生态系统功能与物种多样性呈现单峰曲线关系 ,在区域尺度上为单调上升关系 ;(2 )非生物因素与多样性 -生产力的交互关系吸引了许多实验研究。人们发现 :物种多样性 -生产力关系可能会受到资源供给率和环境扰动的修正 ,环境因素可能是多样性 -生产力关系的幕后操纵者 ;(3)人们开始重视营养级相互作用对于多样性 -生态系统功能关系的影响 ,生态位互补和抽样假说开始被扩展运用到消费者营养级上 ;(4 )人们开始认真思考物种共存机制在多样性 -生态系统功能关系的形成中所扮演的角色。理论模型研究表明 ,不同的物种共存机制会导致不同的多样性 -生产力关系     

Biodiversity and ecosystem productivity: implications for carbon storage

Recent experiments have found that Net Primary Productivity (NPP) can often be a positive saturating function of plant species and functional diversity. These findings raised the possibility that more diverse ecosystems might store more carbon as a result of increased photosynthetic inputs. However, carbon inputs will not only remain in plant biomass, but will be translocated to the soil via root exudation, fine root turnover, and litter fall. Thus, we must consider not just plant productivity (NPP), but also net productivity of the whole ecosystem (NEP), which itself measures net carbon storage. We currently know little about how plant diversity could influence soil processes that return carbon back to the atmosphere, such as heterotrophic respiration and decomposition of organic matter. Nevertheless, it is clear that any effects on such processes could make NPP a poor predictor of whole-ecosystem productivity, and potentially the ability of the ecosystem to store carbon. We examine the range of mechanisms by which plant diversity could influence net ecosystem productivity, incorporating processes involved with carbon uptake (productivity), loss (autotrophic and heterotrophic respiration), and residence time within the system (decomposition rate). Understanding the relationship between plant diversity and ecosystem carbon dynamics must be made a research priority if we wish to provide information relevant to global carbon policy decisions. This goal is entirely feasible if we utilize some basic methods for measuring the major fluxes of carbon into and out of the ecosystem.     

Effects of wind on a shallow lake ecosystem: Resuspension of particles in the Loosdrecht Lakes

Horizontal variation of seston concentration in the shallow, eutrophic Loosdrecht Lakes (A=9.8 km²; =1.9 m) was studied in relation to windspeed and effective fetch. Simple wave theory was applied in order to predict resuspension using wind data from a nearby meteorological station. Most results were consistent with the theory, but a clear limit for the occurrence of resuspension could not be established. Generally, changes in epipelon—the particles at the sediment-water interface — were not directly related to computed frequency of resuspension at the sampling station. The frequency was estimated for 37 grid points over the entire lake. Resuspension was computed to affect high percentages of the lake area in winter. In summer the frequency was much lower, but in June and July 1984 there were days with nearly 50% of the lake area subject to resuspension. The resulting input of particulate organic carbon into the water column during these days was estimated to equal 12–25 times the daytime phytoplankton carbon fixation. Most of the resuspended matter appeared to be redeposited rapidly. The computed frequency of resuspension for the 37 locations of the lake varied between 7 and 48 days in 1984.