Forest biodiversity,ecosystem functioning and the provision of ecosystem services

Forests are critical habitats for biodiversity and they are also essential for the provision of a wide range of ecosystem services that are important to human well-being. There is increasing evidence that biodiversity contributes to forest ecosystem functioning and the provision of ecosystem services. Here we provide a review of forest ecosystem services including biomass production, habitat provisioning services, pollination, seed dispersal, resistance to wind storms, fire regulation and mitigation, pest regulation of native and invading insects, carbon sequestration, and cultural ecosystem services, in relation to forest type, structure and diversity. We also consider relationships between forest biodiversity and multifunctionality, and trade-offs among ecosystem services. We compare the concepts of ecosystem processes, functions and services to clarify their definitions. Our review of published studies indicates a lack of empirical studies that establish quantitative and causal relationships between forest biodiversity and many important ecosystem services. The literature is highly skewed; studies on provisioning of nutrition and energy, and on cultural services, delivered by mixed-species forests are under-represented. Planted forests offer ample opportunity for optimising their composition and diversity because replanting after harvesting is a recurring process. Planting mixed-species forests should be given more consideration as they are likely to provide a wider range of ecosystem services within the forest and for adjacent land uses. This review also serves as the introduction to this special issue of Biodiversity and Conservation on various aspects of forest biodiversity and ecosystem services.     

生物多样性和生态系统功能研究综述

生物多样性和生态系统功能之间关系 ,是生态学和环境科学的热门话题。围绕这一主题 ,文章系统回顾了近 2 0年来的研究历史及学术界的不同观点 ,全面展示了目前在理论和实验领域的主要工作结果和研究进展 ,并对今后的发展趋势和面临的挑战作了展望。理论和实验研究都表明 ,生物多样性趋于与生态系统功能 (稳定性 )呈正相关性 ,但是多样性并非是这种关系的直接驱动力。生态系统功能 (稳定性 )潜在地依赖于物种之间相互作用的强度 ,物种的功能反应特性以及生态系统的类型和尺度等。在生物多样性和生态系统功能的研究中 ,重要的不只是结论 ,还应包括其中所隐含的机制。     

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.     

Ant biodiversity and its relationship to ecosystem functioning: a review

Ants are important components of ecosystems not only because they constitute a great part of the animal biomass but also because they act as ecosystem engineers. Ant biodiversity is incredibly high and these organisms are highly responsive to human impact, which obviously reduces its richness. However, it is not clear how such disturbance damages the maintenance of ant services to the ecosystem. Ants are important in below ground processes through the alteration of the physical and chemical environment and through their effects on plants, microorganisms, and other soil organisms. This review summarizes the information available on ant biodiversity patterns, how it can be quantified, and how biodiversity is affected by human impacts such as land use change, pollution, invasions, and climate change. The role of ants in ecosystems is discussed, mainly from the perspective of the effects of ground-dwelling ants on soil processes and function, emphasizing their role as ecosystem engineers. Some lines of research are suggested after demonstrating the gaps in our current information on ant-soil interactions.     

The relationship between biodiversity and ecosystem functioning in food webs

Recent theoretical and experimental work provides clear evidence that biodiversity loss can have profound impacts on functioning of natural and managed ecosystems and the ability of ecosystems to deliver ecological services to human societies. Work on simplified ecosystems in which the diversity of a single trophic level is manipulated shows that diversity can enhance ecosystem processes such as primary productivity and nutrient retention. Theory also strongly suggests that biodiversity can act as biological insurance against potential disruptions caused by environmental changes. However, these studies generally concern a single trophic level, primary producers for the most part. Changes in biodiversity also affect ecosystem functioning through trophic interactions. Here we review, through the analysis of a simple ecosystem model, several key aspects inherent in multitrophic systems that may strongly affect the relationship between diversity and ecosystem processes. Our analysis shows that trophic interactions have a strong impact on the relationships between diversity and ecosystem functioning, whether the ecosystem property considered is total biomass or temporal variability of biomass at the various trophic levels. In both cases, food-web structure and trade-offs that affect interaction strength have major effects on these relationships. Multitrophic interactions are expected to make biodiversity–ecosystem functioning relationships more complex and non-linear, in contrast to the monotonic changes predicted for simplified systems with a single trophic level.     

The prominence of and biases in biodiversity and ecosystem functioning research

The sub-discipline of biodiversity and ecosystem functioning (BEF) has emerged as a central topic in contemporary ecological research. However, to date no study has evaluated the prominence and publication biases in BEF research. Herein we report the results of a careful quantitative assessment of BEF research published in five core general ecology journals from 1990 to 2007 to determine the position of BEF research within ecology, identify patterns of research effort within BEF research, and discuss their probable proximal and historical causes. The relative importance of BEF publications increased exponentially during the period analyzed and was significantly greater than the average growth of ecological literature, affirming the prominence of BEF as a current paradigm in ecology. However, BEF research exhibited a strong bias toward experimental studies on terrestrial plant communities, with significantly lower effort devoted to the functional aspects of biodiversity in aquatic systems, multiple trophic level systems, and animal or microbial communities. Such trends may be explained by a combination of methodological adequacy and historic epistemological differences in ecological thinking. We suggest that BEF researchers should direct more effort toward the study of aquatic systems and animal communities, emphasize long-term and trophically complex experiments, such as those with multi-trophic microbial communities, employ larger-scale field observational studies and increase the use of integrative and theoretical studies. Many technical and analytical methodologies that are already employed in ecological research, such as stable isotopes, paleobiology, remote sensing, and model selection criteria, can facilitate these aims. Overcoming the above-mentioned shortcomings of current BEF research will greatly improve our ability to predict how biodiversity loss will affect ecosystem processes and services in natural ecosystems.     

Nontrophic interactions, biodiversity, and ecosystem functioning: an interaction web model

Research into the relationship between biodiversity and ecosystem functioning has mainly focused on the effects of species diversity on ecosystem properties in plant communities and, more recently, in food webs. Although there is growing recognition of the significance of nontrophic interactions in ecology, these interactions are still poorly studied theoretically, and their impact on biodiversity and ecosystem functioning is largely unknown. Existing models of mutualism usually consider only one type of species interaction and do not satisfy mass balance constraints. Here, we present a model of an interaction web that includes both trophic and nontrophic interactions and that respects the principle of mass conservation. Nontrophic interactions are represented in the form of interaction modifications. We use this model to study the relationship between biodiversity and ecosystem properties that emerges from the assembly of entire interaction webs. We show that ecosystem properties such as biomass and production depend not only on species diversity but also on species interactions, in particular on the connectance and magnitude of nontrophic interactions, and that the nature, prevalence, and strength of species interactions in turn depend on species diversity. Nontrophic interactions alter the shape of the relationship between biodiversity and biomass and can profoundly influence ecosystem processes.     

Loss of biodiversity and ecosystem functioning in Indo-Malayan peat swamp forests

The tropical peat swamp forests of Indonesia and Malaysia are unusual ecosystems that are rich in endemic species of flora, fauna and microbes despite their extreme acidic, anaerobic, nutrient poor conditions. They are an important refuge for many endangered species including orang utans. Ecosystem functioning is unusual: microbial decomposition is inhibited because the leaves are sclerophyllous and toxic to deter herbivory in the nutrient poor environment, yet bacteria are abundant and active in the surface layers of the peat, where they respire DOC leached from newly fallen leaves. The bacteria are subsequently consumed by aquatic invertebrates that are eaten by fish, and bacterially respired CO₂ is assimilated by algae, so bacteria are thus vital to carbon and nutrient cycling. Peat swamp forests are highly sensitive to the impacts of logging, drainage and fire, due to the interdependence of the vegetation with the peat substrate, which relies on the maintenance of adequate water, canopy cover and leaf litter inputs. Even minor disturbances can increase the likelihood of fire, which is the major cause of CO₂ emissions from regional peat swamp forests and which impact ecosystems worldwide by contributing to climate change. Indo-Malayan peat swamps affect the hydrology of surrounding ecosystems due to their large water storage capacity which slows the passage of floodwaters in wet seasons and maintains stream base flows during dry seasons. These forests are of global importance yet they are inadequately protected and vanishing rapidly, particularly due to agricultural conversion to oil palm, logging, drainage and annual fires.     

Impacts of multiple stressors on biodiversity and ecosystem functioning: the role of species co-tolerance

Ecosystem resistance to a single stressor relies on tolerant species that can compensate for sensitive competitors and maintain ecosystem processes, such as primary production. We hypothesize that resistance to additional stressors depends increasingly on species tolerances being positively correlated (i.e. positive species co-tolerance). Initial exposure to a stressor combined with positive species co-tolerance should reduce the impacts of other stressors, which we term stress-induced community tolerance. In contrast, negative species co-tolerance is expected to result in additional stressors having pronounced additive or synergistic impacts on biologically impoverished functional groups, which we term stress-induced community sensitivity. Therefore, the sign and strength of the correlation between species sensitivities to multiple stressors must be considered when predicting the impacts of global change on ecosystem functioning as mediated by changes in biodiversity.     

Anthropogenic impacts on tropical forest biodiversity: a network structure and ecosystem functioning perspective

Huge areas of diverse tropical forest are lost or degraded every year with dramatic consequences for biodiversity. Deforestation and fragmentation, over-exploitation, invasive species and climate change are the main drivers of tropical forest biodiversity loss. Most studies investigating these threats have focused on changes in species richness or species diversity. However, if we are to understand the absolute and long-term effects of anthropogenic impacts on tropical forests, we should also consider the interactions between species, how those species are organized in networks, and the function that those species perform. I discuss our current knowledge of network structure and ecosystem functioning, highlighting empirical examples of their response to anthropogenic impacts. I consider the future prospects for tropical forest biodiversity, focusing on biodiversity and ecosystem functioning in secondary forest. Finally, I propose directions for future research to help us better understand the effects of anthropogenic impacts on tropical forest biodiversity.     

Density compensation can cause no effect of biodiversity on ecosystem functioning

The role of density compensation (the decline of species density with increasing diversity), in the context of biodiversity and ecosystem functioning, has not been explicitly explored. I used aquatic microbial communities containing bacterivorous consumers (protozoans and rotifers) to investigate whether competition can lead to density compensation and whether density compensation can contribute to the relationship between biodiversity and ecosystem functioning. The experiment employed a nested design in which the consumer diversity gradient (0, 1, 2 or 4 species) was constructed by drawing all possible species or species combinations at each diversity level from a five-species pool. All consumer species coexisted but there was little evidence for overyielding or species dominance, suggesting weak complementarity and sampling effects. Rather, increasing number of consumer species resulted in community-wide density compensation, such that aggregate consumer biomass was unaffected by consumer diversity. Whereas culturable bacterial density declined as consumer diversity increased, total bacterial density showed no discernible response to changes in consumer diversity, a result probably due in part to heterogeneity in bacterial edibility. This study demonstrates the potential for density compensation to shape the relationship between biodiversity and ecosystem functioning.     

Intraspecific traits change biodiversity effects on ecosystem functioning under metal stress

Studies investigating the impacts of biodiversity loss on ecosystem processes have often reached different conclusions, probably because insufficient attention has been paid to some aspects including (1) which biodiversity measure (e.g., species number, species identity or trait) better explains ecosystem functioning, (2) the mechanisms underpinning biodiversity effects, and (3) how can environmental context modulates biodiversity effects. Here, we investigated how species number (one to three species) and traits of aquatic fungal decomposers (by replacement of a functional type from an unpolluted site by another from a metal-polluted site) affect fungal production (biomass acumulation) and plant litter decomposition in the presence and absence of metal stress. To examine the putative mechanisms that explain biodiversity effects, we determined the contribution of each fungal species to the total biomass produced in multicultures by real-time PCR. In the absence of metal, positive diversity effects were observed for fungal production and leaf decomposition as a result of species complementarity. Metal stress decreased diversity effects on leaf decomposition in assemblages containing the functional type from the unpolluted site, probably due to competitive interactions between fungi. However, dominance effect maintained positive diversity effects under metal stress in assemblages containing the functional type from the metal-polluted site. These findings emphasize the importance of intraspecific diversity in modulating diversity effects under metal stress, providing evidence that trait-based diversity measures should be incorporated when examining biodiversity effects.     

Exponential decline of deep-sea ecosystem functioning linked to benthic biodiversity loss

BACKGROUND: Recent investigations suggest that biodiversity loss might impair the functioning and sustainability of ecosystems. Although deep-sea ecosystems are the most extensive on Earth, represent the largest reservoir of biomass, and host a large proportion of undiscovered biodiversity, the data needed to evaluate the consequences of biodiversity loss on the ocean floor are completely lacking. RESULTS: Here, we present a global-scale study based on 116 deep-sea sites that relates benthic biodiversity to several independent indicators of ecosystem functioning and efficiency. We show that deep-sea ecosystem functioning is exponentially related to deep-sea biodiversity and that ecosystem efficiency is also exponentially linked to functional biodiversity. These results suggest that a higher biodiversity supports higher rates of ecosystem processes and an increased efficiency with which these processes are performed. The exponential relationships presented here, being consistent across a wide range of deep-sea ecosystems, suggest that mutually positive functional interactions (ecological facilitation) can be common in the largest biome of our biosphere. CONCLUSIONS: Our results suggest that a biodiversity loss in deep-sea ecosystems might be associated with exponential reductions of their functions. Because the deep sea plays a key role in ecological and biogeochemical processes at a global scale, this study provides scientific evidence that the conservation of deep-sea biodiversity is a priority for a sustainable functioning of the worlds' oceans.     

Global human footprint on the linkage between biodiversity and ecosystem functioning in reef fishes

Difficulties in scaling up theoretical and experimental results have raised controversy over the consequences of biodiversity loss for the functioning of natural ecosystems. Using a global survey of reef fish assemblages, we show that in contrast to previous theoretical and experimental studies, ecosystem functioning (as measured by standing biomass) scales in a non-saturating manner with biodiversity (as measured by species and functional richness) in this ecosystem. Our field study also shows a significant and negative interaction between human population density and biodiversity on ecosystem functioning (i.e., for the same human density there were larger reductions in standing biomass at more diverse reefs). Human effects were found to be related to fishing, coastal development, and land use stressors, and currently affect over 75% of the world's coral reefs. Our results indicate that the consequences of biodiversity loss in coral reefs have been considerably underestimated based on existing knowledge and that reef fish assemblages, particularly the most diverse, are greatly vulnerable to the expansion and intensity of anthropogenic stressors in coastal areas.     

A general multi-trait-based framework for studying the effects of biodiversity on ecosystem functioning

Environmental change is as multifaceted as are the species and communities that respond to these changes. Current theoretical approaches to modeling ecosystem response to environmental change often deal only with single environmental drivers or single species traits, simple ecological interactions, and/or steady states, leading to concern about how accurately these approaches will capture future responses to environmental change in real biological systems. To begin addressing this issue, we generalize a previous trait-based framework to incorporate aspects of frequency dependence, functional complementarity, and the dynamics of systems composed of species that are defined by multiple traits that are tied to multiple environmental drivers. The framework is particularly well suited for analyzing the role of temporal environmental fluctuations in maintaining trait variability and the resultant effects on community response to environmental change. Using this framework, we construct simple models to investigate two ecological problems. First, we show how complementary resource use can significantly enhance the nutrient uptake of plant communities through two different mechanisms related to increased productivity (over-yielding) and larger trait variability. Over-yielding is a hallmark of complementarity and increases the total biomass of the community and, thus, the total rate at which nutrients are consumed. Trait variability also increases due to the lower levels of competition associated with complementarity, thus speeding up the rate at which more efficient species emerge as conditions change. Second, we study systems in which multiple environmental drivers act on species defined by multiple, correlated traits. We show that correlations in these systems can increase trait variability within the community and again lead to faster responses to environmental change. The methodological advances provided here will apply to almost any function that relates species traits and environmental drivers to growth, and should prove useful for studying the effects of climate change on the dynamics of biota.     

On the importance of the negative selection effect for the relationship between biodiversity and ecosystem functioning

Much of our knowledge on biodiversity and ecosystem functioning comes from studies examining the effects of biodiversity on biomass production within a trophic group. A large number of these studies have found that increasing biodiversity tends to increase biomass production, leading many ecologists to believe that there exists a general positive relationship between biodiversity and ecosystem functioning. Here we argue that such a positive relationship may not be general, particularly for ecosystem functions other than biomass. Our argument centers on the potential importance of the negative selection effect, which operates where competitively dominant species do not contribute significantly to the function of interest. We suggest that negative selection effects may be potentially common for non-biomass functions, for which species competitive ability may often be a poor indictor of its functional impact. We conclude that diverse (positive, negative, and neutral) BEF relationships are possible for non-biomass functions and that for a particular function, the exact form of the BEF relationship may depend on how species functional impacts relate to their competitive abilities in the community.     

Tradeoffs and thresholds in the effects of nitrogen addition on biodiversity and ecosystem functioning: evidence from inner Mongolia Grasslands

Nitrogen (N) deposition is widely considered an environmental problem that leads to biodiversity loss and reduced ecosystem resilience; but, N fertilization has also been used as a management tool for enhancing primary production and ground cover, thereby promoting the restoration of degraded lands. However, empirical evaluation of these contrasting impacts is lacking. We tested the dual effects of N enrichment on biodiversity and ecosystem functioning at different organizational levels (i.e., plant species, functional groups, and community) by adding N at 0, 1.75, 5.25, 10.5, 17.5, and 28.0 g N m^?2 yr^?1 for four years in two contrasting field sites in Inner Mongolia: an undisturbed mature grassland and a nearby degraded grassland of the same type. N addition had both quantitatively and qualitatively different effects on the two communities. In the mature community, N addition led to a large reduction in species richness, accompanied by increased dominance of early successional annuals and loss of perennial grasses and forbs at all N input rates. In the degraded community, however, N addition increased the productivity and dominance of perennial rhizomatous grasses, with only a slight reduction in species richness and no significant change in annual abundance. The mature grassland was much more sensitive to N‐induced changes in community structure, likely as a result of higher soil moisture accentuating limitation by N alone. Our findings suggest that the critical threshold for N‐induced species loss to mature Eurasian grasslands is below 1.75 g N m^?2 yr^?1, and that changes in aboveground biomass, species richness, and plant functional group composition to both mature and degraded ecosystems saturate at N addition rates of approximately 10.5 g N m^?2 yr^?1. This work highlights the tradeoffs that exist in assessing the total impact of N deposition on ecosystem function.