Indicators and mechanisms of stability and resilience to climatic and landscape changes in a remnant calcareous grassland

When considering the development of conservation strategies for threatened plant communities it is crucial to understand their resilience to environmental change, taking into account current decline and the occurrence of further habitat fragmentation and climatic changes. Many recent works describe resilience character and elements, but there is little focus on the metrics and indices that describe elements of stability and specific resistance or resilience over the community composition matrix. Communities with strong niche selection might be restricted to specific resistance strategies to cope with environmental changes. This would result in a community at greater risk from increasing fragmentation and climatic changes.In a 35 years survey of relic calcareous grasslands, we looked at measures to identify the resilience mechanisms for stability in the presence and abundance of species. We used techniques of partitioning of temporal beta diversity in nestedness and turnover components, analysis of functional strategy changes and dissimilarity analysis to detect changes in between⿿plot diversity and exchanges.Contrary to expectations, we observe strong resilience with different stabilizing mechanisms both at plot level and exchanges between plots. At the scale of our grassland complex, response diversity and environmental stochasticity allow for the maintenance of high biodiversity under natural perturbations and gradual human-induced environmental changes. This highlights the importance of dispersal, recruitment dynamics and microsite diversity.Community resilience is more than just the sum of species resilience strategies; adaptive management strategies need more emphasis on the variability of conditions, as this can enable or disrupt important community resilience mechanisms.     

Phenotypic diversity and stability of ecosystem processes.

The resistance of an ecosystem to perturbations and the speed at which it recovers after the perturbations, which is called resilience, are two important components of ecosystem stability. It has been suggested that biodiversity increases the resilience and resistance of aggregated ecosystem processes. We test this hypothesis using a theoretical model of a nutrient-limited ecosystem in a heterogeneous environment. We investigate the stability properties of the model for its simplest possible configuration, i.e. , a system consisting of two plant species and their associated detritus and local resource depletion zones. Phenotypic diversity within the plant community is described by differences in the nutrient uptake and mortality rates of the two species. The usual measure of resilience characterizes the system as a whole and thus also applies to aggregated ecosystem processes. As a rule this decreases with increased diversity, though under certain conditions it is maximum for an intermediate value of diversity. Resistance is a property that characterizes each system component and process separately. The resistance of the inorganic nutrient pools, hence of nutrient retention in the ecosystem, decreases with increased diversity. The resistance of both total plant biomass and productivity either monotonically decreases or increases over part of the parameter range with increased diversity. Furthermore, it is very sensitive to parameter values. These results support the view that there is no simple relationship between diversity and stability in equilibrium deterministic systems, whether at the level of populations or aggregated ecosystem processes. We discuss these results in relation to recent experiments.     

Complexity can enhance stability in competitive systems

Empirical observations often indicate that complexity enhances stability, while most theoretical studies, such as May's (1972) classic paper, point to the opposite. Despite the wide generality of these latter theoretical analyses, our examination of the well-known competitive Lotka–Volterra system reveals that increasing complexity (measured in terms of connectance) can enhance species coexistence and persistence in model communities (measured in terms of their feasibility and stability). The high feasibility and stability found for tightly interconnected competitive subsystems might provide an explanation for the clumped structure in food webs.     

Relative ascendency predicts food web robustness

Threats to ecosystems globally from anthropogenic disturbance and climate change requires us to urgently identify the most sensitive biological communities to ensure they are effectively preserved. It is for this reason that understanding and predicting food web stability has been topical within ecology. Food web stability is a multi-faceted concept that represents the ability of a food web to maintain its integrity following disturbance, it includes resistance, resilience and fragility. In this study, we examine the ability of four food web metrics to predict the fragility to random species extinctions in 120 qualitative food webs. We show that three information-based indices out performed food web connectance in predicting fragility, with relative ascendency having the strongest relationship. Relative ascendency was a much stronger predictor of fragility than MacArthur’s stability metric, Average Mutual Information and connectance as it accounted for both the distribution and number of links between species. We also find that most qualitative food webs persist around a central tendency of relative ascendency.     

Species richness facilitates ecosystem resilience in aquatic food webs

1. Many studies indicate that biodiversity in ecosystems affects stability, either by promoting temporal stability of ecosystem attributes or by enhancing ecosystem resistance and resilience to perturbation. The effects on temporal stability are reasonably well understood and documented but effects on resistance and resilience are not. 2. Here, we report results from an aquatic mesocosm experiment in which we manipulated the species richness and composition of aquatic food webs (macrophytes, macro‐herbivores and invertebrate predators), imposed a pulse disturbance (acidification), and monitored the resistance (initial response) and resilience (recovery) of ecosystem productivity and respiration. 3. We found that species‐rich macroinvertebrate communities had higher resilience of whole‐ecosystem respiration, but were not more resistant to perturbations. We also found that resilience and resistance were unaffected by species composition, despite the strong role composition is known to play in determining mean levels of function in these communities. 4. Biodiversity’s effects on resilience were probably mediated through complex pathways affecting phytoplankton and microbial communities (e.g. via changes in nutrient regeneration, grazing or compositional changes) rather than through simpler effects (e.g. insurance effects, enhanced facilitation) although these simpler mechanisms probably played minor roles in enhancing respiration resilience. 5. Current mechanisms for understanding biodiversity’s effects on ecosystem stability have been developed primarily in the context of single‐trophic level communities. These mechanisms may be overly simplistic for understanding the consequences of species richness on ecosystem stability in complex, multi‐trophic food webs where additional factors such as indirect effects and highly variable life‐history traits of species may also be important.     

Diversity enhances community recovery, but not resistance, after drought

1.  There is growing concern that the current loss of biodiversity may negatively affect ecosystem functioning and stability. Although it has been shown that species loss may reduce biomass production and increase temporal variability, experimental evidence that species loss affects ecosystem resistance and resilience after perturbation is limited.
2.  Here, we use the response of experimental plant communities – which differ in diversity – to a natural drought to disentangle the effects of diversity and biomass on resistance, recovery and resilience.
3.  Resistance to drought decreased with diversity, but this pattern was highly dependent upon pre-drought biomass. When corrected for biomass, no relationship between diversity and resistance was observed: at each level of diversity, biomass production was reduced by approximately 30%.
4.  In contrast, recovery (change in biomass production after drought) increased with diversity and was independent of biomass. Resilience (measured as the ratio of post- to pre-drought biomass) was similar at each level of diversity.
5.   Synthesis . On the one hand, our results confirm earlier findings that a positive relationship between diversity and resistance is mainly driven by pre-perturbation performance rather than by diversity. However, the results also show that recovery after drought strongly increased with diversity, independent of performance. We conclude that it is this diversity-dependent recovery which allowed diverse, productive communities to reach the same level of resilience as less diverse (and productive) communities. This finding provides strong experimental evidence for the insurance hypothesis.     

Effects of above-ground plant species composition and diversity on the diversity of soil-borne microorganisms

A coupling of above-ground plant diversity and below-ground microbial diversity has been implied in studies dedicated to assessing the role of macrophyte diversity on the stability, resilience, and functioning of ecosystems. Indeed, above-ground plant communities have long been assumed to drive below-ground microbial diversity, but to date very little is known as to how plant species composition and diversity influence the community composition of micro-organisms in the soil. We examined this relationship in fields subjected to different above-ground biodiversity treatments and in field experiments designed to examine the influence of plant species on soil-borne microbial communities. Culture-independent strategies were applied to examine the role of wild or native plant species composition on bacterial diversity and community structure in bulk soil and in the rhizosphere. In comparing the influence of Cynoglossum officinale (hound's tongue) and Cirsium vulgare (spear thistle) on soil-borne bacterial communities, detectable differences in microbial community structure were confined to the rhizosphere. The colonisation of the rhizosphere of both plants was highly reproducible, and maintained throughout the growing season. In a separate experiment, effects of plant diversity on bacterial community profiles were also only observed for the rhizosphere. Rhizosphere soil from experimental plots with lower macrophyte diversity showed lower diversity, and bacterial diversity was generally lower in the rhizosphere than in bulk soil. These results demonstrate that the level of coupling between above-ground macrophyte communities and below-ground microbial communities is related to the tightness of the interactions involved. Although plant species composition and community structure appear to have little discernible effect on microbial communities inhabiting bulk soil, clear and reproducible changes in microbial community structure and diversity are observed in the rhizosphere. This revised version was published online in August 2006 with corrections to the Cover Date.     

Decline of soil microbial diversity does not influence the resistance and resilience of key soil microbial functional groups following a model disturbance

Analysing the consequences of the decrease in biodiversity for ecosystem functioning and stability has been a major concern in ecology. However, the impact of decline in soil microbial diversity on ecosystem sustainability remains largely unknown. This has been assessed for decomposition, which is insured by a large proportion of the soil microbial community, but not for more specialized and less diverse microbial groups. We determined the impact of a decrease in soil microbial diversity on the stability (i.e. resistance and resilience following disturbance) of two more specialized bacterial functional groups: denitrifiers and nitrite oxidizers. Soil microbial diversity was reduced using serial dilutions of a suspension obtained from a non-sterile soil that led to loss of species with low cell abundance, inoculation of microcosms of the same sterile soil with these serial dilutions, and subsequent incubation to enable establishment of similar cell abundances between treatments. The structure, cell abundance and activity of denitrifying and nitrite-oxidizing communities were characterized after incubation. Increasing dilution led to a progressive decrease in community diversity as assessed by the number of denaturating gradient gel electrophoresis (DGGE) bands, while community functioning was not impaired when cell abundance recovered after incubation. The microcosms were then subjected to a model disturbance: heating to 42 degrees C for 24 h. Abundance, structure and activity of each community were measured 3 h after completion of the disturbance to assess resistance, and after incubation of microcosms for 1 month to assess resilience. Resistance and resilience to the disturbance differed between the two communities, nitrite oxidizers being more affected. However, reducing the diversity of the two microbial functional groups did not impair either their resistance or their resilience following the disturbance. These results demonstrate the low sensitivity of the resistance and resilience of both microbial groups to diversity decline provided that cell abundance is similar between treatments.     

Rhizosphere microbiome functional diversity and pathogen invasion resistance build up during plant development

The rhizosphere microbiome is essential for plant growth and health, and numerous studies have attempted to link microbiome functionality to species and trait composition. However, to date little is known about the actual ecological processes shaping community composition, complicating attempts to steer microbiome functionality. Here, we assess the development of microbial life history and community-level species interaction patterns that emerge during plant development. We use microbial phenotyping to experimentally test the development of niche complementarity and life history traits linked to microbiome performance. We show that the rhizosphere microbiome assembles from pioneer assemblages of species with random resource overlap into high-density, functionally complementary climax communities at later stages. During plant growth, fast-growing species were further replaced by antagonistic and stress-tolerant ones. Using synthetic consortia isolated from different plant growth stages, we demonstrate that the high functional diversity of ‘climax’ microbiomes leads to a better resistance to bacterial pathogen invasion. By demonstrating that different life-history strategies prevail at different plant growth stages and that community-level processes may supersede the importance of single species, we provide a new toolbox to understand microbiome assembly and steer its functionality at a community level.     

Effects of experimentally-enhanced precipitation and nitrogen on resistance,recovery and resilience of a semi-arid grassland after drought

Resistance, recovery and resilience are three important properties of ecological stability, but they have rarely been studied in semi-arid grasslands under global change. We analyzed data from a field experiment conducted in a native grassland in northern China to explore the effects of experimentally enhanced precipitation and N deposition on both absolute and relative measures of community resistance, recovery and resilience—calculated in terms of community cover—after a natural drought. For both absolute and relative measures, communities with precipitation enhancement showed higher resistance and lower recovery, but no change in resilience compared to communities with ambient precipitation in the semi-arid grassland. The manipulated increase in N deposition had little effect on these community stability metrics except for decreased community resistance. The response patterns of these stability metrics to alterations in precipitation and N are generally consistent at community, functional group and species levels. Contrary to our expectations, structural equation modeling revealed that water-driven community resistance and recovery result mainly from changes in community species asynchrony rather than species diversity in the semi-arid grassland. These findings suggest that changes in precipitation regimes may have significant impacts on the response of water-limited ecosystems to drought stress under global change scenarios.     

Complexity of multitrophic interactions in a grassland ecosystem depends on plant species diversity

1.?We studied the theoretical prediction that a loss of plant species richness has a strong impact on community interactions among all trophic levels and tested whether decreased plant species diversity results in a less complex structure and reduced interactions in ecological networks. 2.?Using plant species-specific biomass and arthropod abundance data from experimental grassland plots (Jena Experiment), we constructed multitrophic functional group interaction webs to compare communities based on 4 and 16 plant species. 427 insect and spider species were classified into 13 functional groups. These functional groups represent the nodes of ecological networks. Direct and indirect interactions among them were assessed using partial Mantel tests. Interaction web complexity was quantified using three measures of network structure: connectance, interaction diversity and interaction strength. 3.?Compared with high plant diversity plots, interaction webs based on low plant diversity plots showed reduced complexity in terms of total connectance, interaction diversity and mean interaction strength. Plant diversity effects obviously cascade up the food web and modify interactions across all trophic levels. The strongest effects occurred in interactions between adjacent trophic levels (i.e. predominantly trophic interactions), while significant interactions among plant and carnivore functional groups, as well as horizontal interactions (i.e. interactions between functional groups of the same trophic level), showed rather inconsistent responses and were generally rarer. 4.?Reduced interaction diversity has the potential to decrease and destabilize ecosystem processes. Therefore, we conclude that the loss of basal producer species leads to more simple structured, less and more loosely connected species assemblages, which in turn are very likely to decrease ecosystem functioning, community robustness and tolerance to disturbance. Our results suggest that the functioning of the entire ecological community is critically linked to the diversity of its component plants species.     

Density‐dependent survival varies with species life‐history strategy in a tropical forest

Species coexistence in diverse communities likely results from multiple interacting factors. Mechanisms such as conspecific negative density dependence (CNDD) and varying life‐history strategies related to resource partitioning are known to influence plant fitness, and thereby community composition and diversity. However, we have little understanding of how these mechanisms interact and how they vary across life stages. Here, we document the interaction between CNDD and life‐history strategy, based on growth‐mortality trade‐offs, from seedling to adult tree for 47 species in a tropical forest. Species’ life‐history strategies remained consistent across stages: fast‐growing species had higher mortality than slow‐growing species at all stages. In contrast, mean CNDD was strongest at early life stages (i.e. seedling, sapling). Fast‐growing species tended to suffer greater CNDD than slow‐growing species at several, but not all life stages. Overall, our results demonstrate that coexistence mechanisms interact across multiple life stages to shape diverse tree communities.     

再论生物多样性与生态系统的稳定性

本文在简述生物多样性与生态系统稳定性研究动态的基础上,从生物多样性和稳定性的概念出发,指出忽视多样性和稳定性的生物组织层次可能是造成观点纷争的根源之一。特定生物组织层次的稳定性可能更多地与该层次的多样性特征相关,探讨多样性和稳定性的关系应从不同的生物组织层次上进行,抗动是生态系统多样性与稳定性关系悖论中的重要因子,如果根据扰动的性质,把生态系统（或其他组织层次）区分为受非正常外力干扰和受环境因子时间异质性波动干扰2类系统,稳定性的4个内涵可以理解为：对于受非正常外力干扰的系统而言,抵抗力和恢复力是稳定性适宜的测度指标;对于受环境因子时间异质性波动干扰和系统而言。利用持久性和变异性衡量系统的稳定性则更具实际意义。结合对群落和种群层次多样性与稳定性相关机制的初步讨论,本文认为;在特定的前提下,多样性可以导致稳定性。     

Resilience to fire of phylogenetic diversity across biological domains

Fire alters the structure and composition of above‐ and belowground communities with concurrent shifts in phylogenetic diversity. The inspection of postfire trends in the diversity of ecological communities incorporating phylogenetic information allows to better understand the mechanisms driving fire resilience. While fire reduces plant phylogenetic diversity based on the recruitment of evolutionarily related species with postfire seed persistence, it increases that of soil microbes by limiting soil resources and changing the dominance of competing microbes. Thus, during postfire community reassembly, plant and soil microbes might experience opposing temporal trends in their phylogenetic diversity that are linked through changes in the soil conditions. We tested this hypothesis by investigating the postfire evolution of plant and soil microbial (fungi, bacteria and archaea) communities across three 20‐year chronosequences. Plant phylogenetic diversity increased with time since fire as pioneer seeders facilitate the establishment of distantly related late‐successional shrubs. The postfire increase in plant phylogenetic diversity fostered plant productivity, eventually recovering soil organic matter. These shifts over time in the soil conditions explained the postfire restoration of fungal and bacterial phylogenetic diversity, which decreased to prefire levels, suggesting that evolutionarily related taxa with high relative fitness recover their competitive superiority during community reassembly. The resilience to fire of phylogenetic diversity across biological domains helps preserve the evolutionary history stored in our ecosystems.     

Measuring Functional Diversity in Plant Communities with MixedLife Forms: A Problem of Hardand Soft Attributes

This paper examines the functional relationships among species in an Australian rangeland community with mixed life forms. It follows a previous study (Walker and others 1999) that explored the role of dominant and minor species in maintaining functional diversity and resilience in a rangeland ecosystem. Unlike our previous results, which were based on estimates of five plant functional attributes, the dominant species in this second community apparently are functionally no more dissimilar to each other than to all other species. We suggest that the lack of clear results in mixed life form communities represents a confounding of the relationships between the “hard” attributes that actually govern how a plant performs in an ecosystem and the “soft” attributes that we use as surrogates. There are very few data on the hard functional attributes of plant species and consequently little information on precisely how the (soft) measurable traits are related to their imputed functions. What evidence there is shows that although the relationships are strong within life forms, they differ between life forms. This poses a problem for the development of research relating plant biodiversity to ecosystem function. Until such a database is developed, it will be very difficult to advance our understanding and measurement of functional diversity in mixed life form communities. Received 30 April 2001; Accepted 23 January 2002.     

土壤微生物群落抵抗力和恢复力研究进展

人类活动的不断加剧使得土壤生态系统承受着环境干扰压力。土壤微生物受到环境干扰的响应程度（抵抗力）及恢复至原来状态的能力（恢复力）决定着土壤生态系统的可持续性。梳理和总结了土壤微生物群落对环境干扰的抵抗力和恢复力方面的研究进展。首先,在介绍土壤微生物群落抵抗力和恢复力概念的基础上,阐述了通过评估微生物群落的结构和功能的变化来系统表征抵抗力和恢复力;随后,分析了最近十年（2012-2021年）有关文献,发现土壤微生物群落的结构和（或）功能在环境干扰后的恢复力总体较弱,但耕作、有机物料添加和轮作等农田管理措施下的响应趋势表现出一定的规律性;继而,从个体水平的休眠和胁迫忍耐、种群水平的生存策略、群落水平的多样性和相互作用以及生态系统水平的历史遗留效应等方面分析了土壤微生物群落抵抗力和恢复力的维持机制;最后,从功能性状、多功能性和植物-土壤微生物整体性对未来研究做出了展望,以期为构建土壤健康评价体系及预测环境干扰对土壤功能的影响提供科学依据。     

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.     

Biodiversity Effects on Biomass Production and Invasion Resistance in Annual versus Perennial Plant Communities

In a field experiment we constructed two different communities using both annual and perennial plant species, in which species diversity is experimentally manipulated. We want to test the relationships between species diversity and biomass production and invasibility and the possible mechanisms driving this relationships, especially, whether the identical mechanisms drive both diversity-production and diversity-invasibility relationships. Our results indicated that a positive diversity-production relationship and negative diversity-invasibility and production-invasibility relationships emerged in two different communities. However, the mechanisms underlying are different in two communities. In the annual communities, the observed positive diversity-production and negative diversity-invasibility relationships are linked by the sampling effect. In the perennial communities, however, the mechanism responsible for these observed relationships are the complementarity effect. Our results also found that, in addition to species diversity, species composition also play an important role in governing the observed relationship. The results of our study suggest that because species in different communities may differ in their life history, biological and physiological traits, mechanisms responsible for the observed relationship between diversity and biomass production and invasibiltiy are likely different.     

Differentiating between effects of invasion and diversity: impacts of aboveground plant communities on belowground fungal communities

Exotic plant species can affect soil microbial communities with the potential for community and ecosystem feedbacks. Yet, separating the effects of exotics from confounded changes in plant community diversity still remains a challenge. We focused on how plant diversity and native or exotic life history affected root fungi because of their significant roles in community and ecosystem processes. Specifically, we examined how fungi colonizing plant roots were affected by plant richness (one, two or four species) replicated across a range of plant community mixtures (natives, exotics, native-exotic mixtures). Fungal biomass inside roots was affected independently by plant richness and mixture, while root fungal community composition was affected only by plant richness. Extraradical networks also increased in size with plant richness. By contrast, plant biomass was a function of plant mixture, with natives consistently smaller than exotics and native-exotic mixtures intermediate. Plant invasions may have an impact on the belowground community primarily through their effects on diversity, at least in the short-term. Disentangling the effects of diversity and invasion on belowground microbial communities can help us to understand both the controllers of belowground resilience and mechanisms of successful colonization and spread of exotic plants.