How are biodiversity and carbon stock recovered during tropical forest restoration? Supporting the ecological paradigms and political context involved

To meet agendas for biodiversity conservation and mitigation of climate change, large-scale restoration initiatives propose ecological restoration as an alternative that can reconcile these two objectives. In ongoing ecosystem restoration, increased diversity is always associated with increased productivity (and consequent carbon stock), which is among the most important ecosystem functions. The ecological paradigm of this association is that ecosystem biodiversity (B) is positively related to both ecosystem functions and services (EF and ES). However, BEF and BES relationships vary spatially and temporally, which makes understanding these relationships relevant and important for practical restoration actions. In this study, we asked how biodiversity and carbon stock recovery occurs during tropical forest restoration. We reviewed literature of the relationships between BEF and BES in the context of ecological restoration and asked whether ecological restoration can recover both. In addition, we conducted a metadata analysis of studies on the recovery of biodiversity and biomass in regenerating tropical forests (n = 83) to find the best model that describes this relationship. In general, studies showed that ecosystem biodiversity and productivity are positively related, and that restoration can recover both. We found an asymptotic and positive correlation between biodiversity and biomass in tropical forests, suggesting limitation of the mutual gains of these two ecosystem properties during restoration. We discuss these results in the context of ecological theory and the practice of ecological restoration.     

生物多样性与生态系统服务——关系、权衡与管理

生物多样性和生态系统服务是人类生存和社会经济可持续发展的物质基础,应对生物多样性丧失和生态系统服务退化问题已经成为继气候变化之后的又一个全球性环境热点问题。生物多样性是生态系统生产力、稳定性、抵抗生物入侵以及养分动态的主要决定因素,生物多样性越高,生态系统功能性状的范围越广,生态系统服务质量就越高越稳定。目的是探讨生物多样性和生态系统服务之间的关系:(1)明确了生物多样性与生态系统过程、功能、服务之间的关系;(2)生物多样性在生态系统服务中的角色:生物多样性在不同的空间尺度通过各种形式的运行机制与生态系统服务产生联系,生物多样性是生态系统过程的调节者,是巩固生态系统服务的一个重要因素,生物多样性也是一种终极的生态系统服务,并在遗传和物种水平上直接贡献了其利益和价值;(3)生物多样性与生态系统服务权衡和协同关系的研究可以更好的帮助管理人员做出有利的决策和保护工作,也是制定规划和适应策略以减少生物多样性危机带来的不利影响的基础;(4)生物多样性与生态系统服务的关系在不同的时间和空间尺度上是不恒定的,有必要共同确定生态系统服务和生物多样性的空间格局,以有效和可持续的进行生态系统管理;(5)虽然生物多样性保护和生态系统管理还存在许多不确定性,但相关理论应该在管理、保护和恢复生态系统中发挥重要作用。研究提出了进一步研究的领域,以促进生物多样性保护和生态系统服务提供之间的协同作用。希望对相关领域的研究有所帮助。     

How community adaptation affects biodiversity–ecosystem functioning relationships

Evidence is growing that evolutionary dynamics can impact biodiversity–ecosystem functioning (BEF) relationships. However the nature of such impacts remains poorly understood. Here we use a modelling approach to compare random communities, with no trait evolutionary fine‐tuning, and co‐adapted communities, where traits have co‐evolved, in terms of emerging biodiversity–productivity, biodiversity–stability and biodiversity–invasion relationships. Community adaptation impacted most BEF relationships, sometimes inverting the slope of the relationship compared to random communities. Biodiversity–productivity relationships were generally less positive among co‐adapted communities, with reduced contribution of sampling effects. The effect of community‐adaptation, though modest regarding invasion resistance, was striking regarding invasion tolerance: co‐adapted communities could remain very tolerant to invasions even at high diversity. BEF relationships are thus contingent on the history of ecosystems and their degree of community adaptation. Short‐term experiments and observations following recent changes may not be safely extrapolated into the future, once eco‐evolutionary feedbacks have taken place.     

What does biodiversity actually do? A review for managers and policy makers

Conservation managers and policy makers must often justify the need for protection of biodiversity. However, results of scientific studies testing for a positive value of biodiversity in terms of community stability and ecosystem function have been complex and inconsistent. We review recent information on the consequences of loss of biodiversity for natural systems. The relationships described vary with scale of interest – for instance, biodiversity at local scales typically has strong effects on ecosystem function, although the opposite relationship is often found at regional scales. These inconsistencies lead to some concern as to whether these relationships can be used to justify biodiversity protection. This is particularly relevant to policy, where holistic protection of biodiversity has most often been mooted and justified. For managers, who most often work to protect single species, communities or ecosystem functions, biodiversity research has failed to address questions of critical concern such as consequences of the loss of rare species or the identification of functional keystone species. For the general public, we believe that the confusion and debate surrounding biodiversity and ecosystem function relationships is in danger of eroding the positive value society places on biodiversity. We further warn that using those relationships in policy documents as justifications for biodiversity protection is fraught with difficulties. Finally, we contend that biodiversity research has largely not addressed issues of concern to conservation managers, and list a set of priorities for relevant research on the consequences of biodiversity loss.     

Unanticipated impacts of spatial variance of biodiversity on plant productivity

Experiments on biodiversity have shown that productivity is often a decelerating monotonic function of biodiversity. A property of nonlinear functions, known as Jensen's inequality, predicts negative effects of the variance of predictor variables on the mean of response variables. One implication of this relationship is that an increase in spatial variability of biodiversity can cause dramatic decreases in the mean productivity of the system. Here I quantify these effects by conducting a meta‐analysis of experimental data on biodiversity–productivity relationships in grasslands and using the empirically derived estimates of parameters to simulate various scenarios of levels of spatial variance and mean values of biodiversity. Jensen's inequality was estimated independently using Monte Carlo simulations and quadratic approximations. The median values of Jensen's inequality estimated with the first method ranged from 3.2 to 26.7%, whilst values obtained with the second method ranged from 5.0 to 45.0%. Meta‐analyses conducted separately for each combination of simulated values of mean and spatial variance of biodiversity indicated that effect sizes were significantly larger than zero in all cases. Because patterns of biodiversity are becoming increasingly variable under intense anthropogenic pressure, the impact of loss of biodiversity on productivity may be larger than current estimates indicate.     

Biodiversity Increases the Productivity and Stability of Phytoplankton Communities

Global biodiversity losses provide an immediate impetus to elucidate the relationships between biodiversity, productivity and stability. In this study, we quantified the effects of species richness and species combination on the productivity and stability of phytoplankton communities subject to predation by a single rotifer species. We also tested one mechanism of the insurance hypothesis: whether large, slow-growing, potentially-defended cells would compensate for the loss of small, fast-growing, poorly-defended cells after predation. There were significant effects of species richness and species combination on the productivity, relative yield, and stability of phytoplankton cultures, but the relative importance of species richness and combination varied with the response variables. Species combination drove patterns of productivity, whereas species richness was more important for stability. Polycultures containing the most productive single species, Dunaliella, were consistently the most productive. Yet, the most species rich cultures were the most stable, having low temporal variability in measures of biomass. Polycultures recovered from short-term negative grazing effects, but this recovery was not due to the compensation of large, slow-growing cells for the loss of small, fast-growing cells. Instead, polyculture recovery was the result of reduced rotifer grazing rates and persisting small species within the polycultures. Therefore, although an insurance effect in polycultures was found, this effect was indirect and unrelated to grazing tolerance. We hypothesize that diverse phytoplankton assemblages interfered with efficient rotifer grazing and that this “interference effect” facilitated the recovery of the most productive species, Dunaliella. In summary, we demonstrate that both species composition and species richness are important in driving patterns of productivity and stability, respectively, and that stability in biodiverse communities can result from an alteration in consumer functioning. Our findings underscore the importance of predator-prey dynamics in determining the relationships between biodiversity, productivity and stability in producer communities.     

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。     

Herbivore impact on grassland plant diversity depends on habitat productivity and herbivore size

Mammalian herbivores can have pronounced effects on plant diversity but are currently declining in many productive ecosystems through direct extirpation, habitat loss and fragmentation, while being simultaneously introduced as livestock in other, often unproductive, ecosystems that lacked such species during recent evolutionary times. The biodiversity consequences of these changes are still poorly understood. We experimentally separated the effects of primary productivity and herbivores of different body size on plant species richness across a 10-fold productivity gradient using a 7-year field experiment at seven grassland sites in North America and Europe. We show that assemblages including large herbivores increased plant diversity at higher productivity but decreased diversity at low productivity, while small herbivores did not have consistent effects along the productivity gradient. The recognition of these large-scale, cross-site patterns in herbivore effects is important for the development of appropriate biodiversity conservation strategies.     

Will a large complex system be productive?

While the relationship between food web complexity and stability has been well documented, how complexity affects productivity remains elusive. In this study, we combine food web theory and a data set of 149 aquatic food webs to investigate the effect of complexity (i.e. species richness, connectance, and average interaction strength) on ecosystem productivity. We find that more complex ecosystems tend to be more productive, although different facets of complexity have contrasting effects. A higher species richness and/or average interaction strength increases productivity, whereas a higher connectance often decreases it. These patterns hold not only between realized complexity and productivity, but also characterize responses of productivity to simulated declines of complexity. Our model also predicts a negative association between productivity and stability along gradients of complexity. Empirical analyses support our predictions on positive complexity-productivity relationships and negative productivity-stability relationships. Our study provides a step forward towards reconciling ecosystem complexity, productivity and stability.     

Animal and plant space-use drive plant diversity–productivity relationships

Plant community productivity generally increases with biodiversity, but the strength of this relationship exhibits strong empirical variation. In meta-food-web simulations, we addressed if the spatial overlap in plants' resource access and animal space-use can explain such variability. We found that spatial overlap of plant resource access is a prerequisite for positive diversity–productivity relationships, but causes exploitative competition that can lead to competitive exclusion. Space-use of herbivores causes apparent competition among plants, resulting in negative relationships. However, space-use of larger top predators integrates sub-food webs composed of smaller species, offsetting the negative effects of exploitative and apparent competition and leading to strongly positive diversity–productivity relationships. Overall, our results show that spatial overlap of plants' resource access and animal space-use can greatly alter the strength and sign of such relationships. In particular, the scaling of animal space-use effects opens new perspectives for linking landscape processes without effects on biodiversity to productivity patterns.     

Primary productivity and species richness: relationships among functional guilds, residency groups and vagility classes at multiple spatial scales

One of the major determinants of species richness is the amount of energy available, often measured as primary productivity. Heterogeneity of environmental variables has also been found to influence species richness. Predicting species distributions across landscapes and identifying areas that have high species richness, or vulnerable groups of species, is useful for land management. Remotely sensed data may help identify such areas, with the Normalized Difference Vegetation Index (NDVI) providing an estimate of primary productivity. We examined the relationship between maximum productivity (NDVI), heterogeneity of productivity, and species richness of birds and butterflies at multiple spatial scales. We also explored relationships between productivity, functional guilds and residency groups of birds, and vagility classes of butterflies. Positive linear relationships between maximum NDVI and number of functional guilds of birds were found at two spatial scales. We also found positive linear relationships between maximum NDVI and species richness of neotropical migrant birds at two scales. Heterogeneity of NDVI, by contrast, was negatively associated with number of functional guilds of birds and species richness of resident birds. Maximum NDVI was associated with species richness of all butterflies and of the most vagile butterflies. No association was found between heterogeneity of NDVI and species richness of butterflies. In the Great Basin, where high greenness and availability of water correspond to areas of high species richness and maximum NDVI, our results suggest that NDVI can provide a reliable basis for stratifying surveys of biodiversity, by highlighting areas of potentially high biodiversity across large areas. Measures of heterogeneity of NDVI appear to be less useful in explaining species richness.     

Understanding diversity–stability relationships: towards a unified model of portfolio effects

A major ecosystem effect of biodiversity is to stabilise assemblages that perform particular functions. However, diversity–stability relationships (DSRs) are analysed using a variety of different population and community properties, most of which are adopted from theory that makes several restrictive assumptions that are unlikely to be reflected in nature. Here, we construct a simple synthesis and generalisation of previous theory for the DSR. We show that community stability is a product of two quantities: the synchrony of population fluctuations, and an average species‐level population stability that is weighted by relative abundance. Weighted average population stability can be decomposed to consider effects of the mean‐variance scaling of abundance, changes in mean abundance with diversity and differences in species' mean abundance in monoculture. Our framework makes explicit how unevenness in the abundances of species in real communities influences the DSR, which occurs both through effects on community synchrony, and effects on weighted average population variability. This theory provides a more robust framework for analysing the results of empirical studies of the DSR, and facilitates the integration of findings from real and model communities.     

Biodiversity, productivity and the temporal stability of productivity: patterns and processes

Theory predicts that the temporal stability of productivity, measured as the ratio of the mean to the standard deviation of community biomass, increases with species richness and evenness. We used experimental species mixtures of grassland plants to test this hypothesis and identified the mechanisms involved. Additionally, we tested whether biodiversity, productivity and temporal stability were similarly influenced by particular types of species interactions. We found that productivity was less variable among years in plots planted with more species. Temporal stability did not depend on whether the species were planted equally abundant (high evenness) or not (realistically low evenness). Greater richness increased temporal stability by increasing overyielding, asynchrony of species fluctuations and statistical averaging. Species interactions that favoured unproductive species increased both biodiversity and temporal stability. Species interactions that resulted in niche partitioning or facilitation increased both productivity and temporal stability. Thus, species interactions can promote biodiversity and ecosystem services.     

What determines the relationship between plant diversity and habitat productivity?

The relationship between biodiversity and habitat productivity has been a fundamental topic in ecology. Although the relationship between these parameters may exhibit different shapes, the unimodal shape has been frequently encountered. The decrease in diversity at high productivity has usually been attributed to competitive exclusion. We suggest that evolutionary history and dispersal limitation may be even more important in shaping the diversity–productivity relationship. On a global scale, unimodal diversity–productivity relationships dominate in temperate regions, whereas positive relationships are more common in the tropics. This difference can be accounted for by contrasting evolutionary history. Temperate regions have smaller species pools for productive habitats since these habitats have been scarce historically for speciation, while the opposite is true for the tropics. In addition, dispersal within a region may limit diversity either due to the lack of dispersal syndromes at low productivity or the low number of diaspores at high productivity. Thereafter, biotic interactions (competition and facilitation) can shape the relationship. All these processes can act independently or concurrently. We recommend that the common approach to examining empirical diversity–environmental relationships should start with the role of large‐scale processes such as evolutionary history and dispersal limitation, followed by influences associated with ecological interactions.     

Temporal stability in forest productivity increases with tree diversity due to asynchrony in species dynamics

Theory predicts a positive relationship between biodiversity and stability in ecosystem properties, while diversity is expected to have a negative impact on stability at the species level. We used virtual experiments based on a dynamic simulation model to test for the diversity–stability relationship and its underlying mechanisms in Central European forests. First our results show that variability in productivity between stands differing in species composition decreases as species richness and functional diversity increase. Second we show temporal stability increases with increasing diversity due to compensatory dynamics across species, supporting the biodiversity insurance hypothesis. We demonstrate that this pattern is mainly driven by the asynchrony of species responses to small disturbances rather than to environmental fluctuations, and is only weakly affected by the net biodiversity effect on productivity. Furthermore, our results suggest that compensatory dynamics between species may enhance ecosystem stability through an optimisation of canopy occupancy by coexisting species.     

Bacterial diversity patterns along a gradient of primary productivity

Primary productivity is a key determinant of biodiversity patterns in plants and animals but has not previously been shown to affect bacterial diversity. We examined the relationship between productivity and bacterial richness in aquatic mesocosms designed to mimic small ponds. We observed that productivity could influence the composition and richness of bacterial communities. We showed that, even within the same system, different bacterial taxonomic groups could exhibit different responses to changes in productivity. The richness of members of the Cytophaga‐Flavobacteria‐Bacteroides group exhibited a significant hump‐shaped relationship with productivity, as is often observed for plant and animal richness in aquatic systems. In contrast, we observed a significant U‐shaped relationship between richness and productivity for α‐proteobacteria and no discernable relationship for β‐proteobacteria. We show, for the first time, that bacterial diversity varies along a gradient of primary productivity and thus make an important step towards understanding processes responsible for the maintenance of bacterial biodiversity.     

Non-neutral community dynamics: empirical predictions for ecosystem function and diversity from linearized consumer–resource interactions

A general model of linearized species interactions, essentially Lotka–Volterra theory, applied to questions of biodiversity has previously been shown to be a powerful tool for understanding local species–abundance patterns and community responses to environmental change for a single trophic level. Here this approach is extended to predict community composition and responses to environmental changes in trophically structured systems. We show how resource and consumer species richness and their relative abundances vary with the means and variances in enrichment level and strengths of intra- and interspecific interactions. Also demonstrated are the responses of local resource and consumer species richness to the global species pools at both trophic levels, as well as the covariation with net resource productivity. These predictions for resource and consumer specific responses to changes in environmental enrichment and global biodiversity are directly testable.     

Grazing weakens temporal stabilizing effects of diversity in the Eurasian steppe

Many biodiversity experiments have demonstrated that plant diversity can stabilize productivity in experimental grasslands. However, less is known about how diversity–stability relationships are mediated by grazing. Grazing is known for causing species losses, but its effects on plant functional groups (PFGs) composition and species asynchrony, which are closely correlated with ecosystem stability, remain unclear. We conducted a six‐year grazing experiment in a semi‐arid steppe, using seven levels of grazing intensity (0, 1.5, 3.0, 4.5, 6.0, 7.5, and 9.0 sheep per hectare) and two grazing systems (i.e., a traditional, continuous grazing system during the growing period (TGS), and a mixed one rotating grazing and mowing annually (MGS)), to examine the effects of grazing system and grazing intensity on the abundance and composition of PFGs and diversity–stability relationships. Ecosystem stability was similar between mixed and continuous grazing treatments. However, within the two grazing systems, stability was maintained through different pathways, that is, along with grazing intensity, persistence biomass variations in MGS, and compensatory interactions of PFGs in their biomass variations in TGS. Ecosystem temporal stability was not decreased by species loss but rather remain unchanged by the strong compensatory effects between PFGs, or a higher grazing‐induced decrease in species asynchrony at higher diversity, and a higher grazing‐induced increase in the temporal variation of productivity in diverse communities. Ecosystem stability of aboveground net primary production was not related to species richness in both grazing systems. High grazing intensity weakened the temporal stabilizing effects of diversity in this semi‐arid grassland. Our results demonstrate that the productivity of dominant PFGs is more important than species richness for maximizing stability in this system. This study distinguishes grazing intensity and grazing system from diversity effects on the temporal stability, highlighting the need to better understand how grazing regulates ecosystem stability, plant diversity, and their synergic relationships.     

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.     

Trade-offs and synergies between ecosystem productivity and stability in temperate grasslands

Aim

It is crucial to monitor how the productivity of grasslands varies with its temporal stability for management of these ecosystems. However, identifying the direction of the productivity–stability relationship remains challenging because ecological stability has multiple components that can display neutral, positive or negative covariations. Furthermore, evidence suggests that the direction of the productivity–stability relationship depends on the biotic interactions and abiotic conditions that underlie ecosystem productivity and stability. We decipher the relationships between grassland productivity and two components of its stability in four habitat types with contrasting environments and flora.

Location

France.

Time period

2000–2020.

Major taxa

Grassland plant species.

Methods

We used c. 20,000 vegetation plots spread across French permanent grasslands and remotely sensed vegetation indices to quantify grassland productivity and temporal stability. We decomposed stability into constancy (i.e., temporal invariability) and resistance (i.e., maximum deviation from average) and deciphered the direct and indirect effects of abiotic (namely growing season length and nitrogen input) and biotic (namely plant taxonomic diversity, trait diversity and community-weighted mean traits) factors on productivity–stability relationships using structural equation models.

Results

We found a positive relationship between productivity and constancy and a negative relationship between productivity and resistance in all habitats. Abiotic factors had stronger effects on productivity and stability compared with biotic factors. A longer growing season enhanced grassland productivity and constancy. Nitrogen input had positive and negative effects on grassland productivity and resistance, respectively. Trait values affected the constancy and resistance of grassland more than taxonomic and trait diversity, with effects varying from one habitat to another. Productivity was not related to any biotic factor.

Main conclusions

Our findings reveal how vital it is to consider both the multiple components of stability and the interaction between environment and biodiversity to gain an understanding of the relationships between productivity and stability in real-world ecosystems, which is a crucial step for sustainable grassland management.