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.     

Resilience and stability of a pelagic marine ecosystem

The accelerating loss of biodiversity and ecosystem services worldwide has accentuated a long-standing debate on the role of diversity in stabilizing ecological communities and has given rise to a field of research on biodiversity and ecosystem functioning (BEF). Although broad consensus has been reached regarding the positive BEF relationship, a number of important challenges remain unanswered. These primarily concern the underlying mechanisms by which diversity increases resilience and community stability, particularly the relative importance of statistical averaging and functional complementarity. Our understanding of these mechanisms relies heavily on theoretical and experimental studies, yet the degree to which theory adequately explains the dynamics and stability of natural ecosystems is largely unknown, especially in marine ecosystems. Using modelling and a unique 60-year dataset covering multiple trophic levels, we show that the pronounced multi-decadal variability of the Southern California Current System (SCCS) does not represent fundamental changes in ecosystem functioning, but a linear response to key environmental drivers channelled through bottom-up and physical control. Furthermore, we show strong temporal asynchrony between key species or functional groups within multiple trophic levels caused by opposite responses to these drivers. We argue that functional complementarity is the primary mechanism reducing community variability and promoting resilience and stability in the SCCS.     

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。     

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。     

Community stability,complexity and species life history strategies

The essence of the contradiction between traditional ecological complexity-stability hypothesis and recent theoretical results is clarified. The distinction between resilience and resistance is stressed. The possibilities of field verification of May's model are discussed. No satisfactory method for estimation of connectance and mean interaction strength in plant communities has been found. Relation between these parameters and stability in real communities remains an open question. The relation between connectance and stability (resilience) in purely competitive model communities is more complicated than May's rule predicts. The certain value of connectance having been achieved, stability increases with increasing connectance. We assessed the positive relation between species diversity and resistance, and negative relation between species diversity and resilience in plant communities during old-field succession in xeric habitat. But there is no causal relationship between species diversity and both kinds of stability. Resistance and resilience of the plant communities studied were determined primarily by life history strategies of constituent species. The results are interpreted in terms of Grimes' life history strategies and imply the validity of Gleasonian, population-centered explanation of community phenomena.The nomenclature of plants follows F. Ehrendorfer, 1973, Liste der Gefässpflanzen Mitteleuropas, 2. Aufl. G. Fischer, Stuttgart.Acknowledgements. We thank Pavel Kindlmann for helpful discussion. We are also grateful to reviewers for suggesting ways to improve this paper.     

Response diversity determines the resilience of ecosystems to environmental change

A growing body of evidence highlights the importance of biodiversity for ecosystem stability and the maintenance of optimal ecosystem functionality. Conservation measures are thus essential to safeguard the ecosystem services that biodiversity provides and human society needs. Current anthropogenic threats may lead to detrimental (and perhaps irreversible) ecosystem degradation, providing strong motivation to evaluate the response of ecological communities to various anthropogenic pressures. In particular, ecosystem functions that sustain key ecosystem services should be identified and prioritized for conservation action. Traditional diversity measures (e.g. ‘species richness’) may not adequately capture the aspects of biodiversity most relevant to ecosystem stability and functionality, but several new concepts may be more appropriate. These include ‘response diversity’, describing the variation of responses to environmental change among species of a particular community. Response diversity may also be a key determinant of ecosystem resilience in the face of anthropogenic pressures and environmental uncertainty. However, current understanding of response diversity is poor, and we see an urgent need to disentangle the conceptual strands that pervade studies of the relationship between biodiversity and ecosystem functioning. Our review clarifies the links between response diversity and the maintenance of ecosystem functionality by focusing on the insurance hypothesis of biodiversity and the concept of functional redundancy. We provide a conceptual model to describe how loss of response diversity may cause ecosystem degradation through decreased ecosystem resilience. We explicitly explain how response diversity contributes to functional compensation and to spatio‐temporal complementarity among species, leading to long‐term maintenance of ecosystem multifunctionality. Recent quantitative studies suggest that traditional diversity measures may often be uncoupled from measures (such as response diversity) that may be more effective proxies for ecosystem stability and resilience. Certain conclusions and recommendations of earlier studies using these traditional measures as indicators of ecosystem resilience thus may be suspect. We believe that functional ecology perspectives incorporating the effects and responses of diversity are essential for development of management strategies to safeguard (and restore) optimal ecosystem functionality (especially multifunctionality). Our review highlights these issues and we envision our work generating debate around the relationship between biodiversity and ecosystem functionality, and leading to improved conservation priorities and biodiversity management practices that maximize ecosystem resilience in the face of uncertain environmental change.     

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.     

A Diverse Assemblage of Reef Corals Thriving in a Dynamic Intertidal Reef Setting (Bonaparte Archipelago,Kimberley, Australia)

The susceptibility of reef-building corals to climatic anomalies is well documented and a cause of great concern for the future of coral reefs. Reef corals are normally considered to tolerate only a narrow range of climatic conditions with only a small number of species considered heat-tolerant. Occasionally however, corals can be seen thriving in unusually harsh reef settings and these are cause for some optimism about the future of coral reefs. Here we document for the first time a diverse assemblage of 225 species of hard corals occurring in the intertidal zone of the Bonaparte Archipelago, north western Australia. We compare the environmental conditions at our study site (tidal regime, SST and level of turbidity) with those experienced at four other more typical tropical reef locations with similar levels of diversity. Physical extremes in the Bonaparte Archipelago include tidal oscillations of up to 8 m, long subaerial exposure times (>3.5 hrs), prolonged exposure to high SST and fluctuating turbidity levels. We conclude the timing of low tide in the coolest parts of the day ameliorates the severity of subaerial exposure, and the combination of strong currents and a naturally high sediment regime helps to offset light and heat stress. The low level of anthropogenic impact and proximity to the Indo-west Pacific centre of diversity are likely to further promote resistance and resilience in this community. This assemblage provides an indication of what corals may have existed in other nearshore locations in the past prior to widespread coastal development, eutrophication, coral predator and disease outbreaks and coral bleaching events. Our results call for a re-evaluation of what conditions are optimal for coral survival, and the Bonaparte intertidal community presents an ideal model system for exploring how species resilience is conferred in the absence of confounding factors such as pollution.     

Species and genotype diversity drive community and ecosystem properties in experimental microcosms

Species diversity is important to ecosystems because of the increased probability of including species that are strong interactors and/or because multiple-species communities are more efficient at using resources due to synergisms and resource partitioning. Genetic diversity also contributes to ecosystem function through effects on primary productivity, community structure and resilience, and modulating energy and nutrient fluxes. Lacking are studies investigating the relationship between ecosystem function and diversity where hierarchical levels of biological diversity are systematically varied during experimentation. In this experiment, we manipulated both species and genotypic diversity of two Daphnia species in microcosms initially seeded with Chlamydomonas and measured community- and ecosystem-level properties to determine which level of diversity was most important for explaining variation in the property. Our results show that species diversity alters bacterial community composition while high genotypic diversity reduces bacterial richness and primary productivity. In addition, the highest levels of genotypic and species richness appear to increase community and ecosystem stability. These findings reveal that species and genotypic diversity are significant drivers of community and ecosystem properties and stability.     

Spatial Heterogeneity Regulates Plant-Pollinator Networks across Multiple Landscape Scales

Mutualistic plant-pollinator interactions play a key role in biodiversity conservation and ecosystem functioning. In a community, the combination of these interactions can generate emergent properties, e.g., robustness and resilience to disturbances such as fluctuations in populations and extinctions. Given that these systems are hierarchical and complex, environmental changes must have multiple levels of influence. In addition, changes in habitat quality and in the landscape structure are important threats to plants, pollinators and their interactions. However, despite the importance of these phenomena for the understanding of biological systems, as well as for conservation and management strategies, few studies have empirically evaluated these effects at the network level. Therefore, the objective of this study was to investigate the influence of local conditions and landscape structure at multiple scales on the characteristics of plant-pollinator networks. This study was conducted in agri-natural lands in Chapada Diamantina, Bahia, Brazil. Pollinators were collected in 27 sampling units distributed orthogonally along a gradient of proportion of agriculture and landscape diversity. The Akaike information criterion was used to select models that best fit the metrics for network characteristics, comparing four hypotheses represented by a set of a priori candidate models with specific combinations of the proportion of agriculture, the average shape of the landscape elements, the diversity of the landscape and the structure of local vegetation. The results indicate that a reduction of habitat quality and landscape heterogeneity can cause species loss and decrease of networks nestedness. These structural changes can reduce robustness and resilience of plant-pollinator networks what compromises the reproductive success of plants, the maintenance of biodiversity and the pollination service stability. We also discuss the possible explanations for these relationships and the implications for landscape planning in agricultural areas.     

Taxonomical vs. functional responses of bee communities to fire in two contrasting climatic regions

1. Valuable insights into mechanisms of community responses to environmental change can be gained by analysing in tandem the variation in functional and taxonomic composition along environmental gradients. 2. We assess the changes in species and functional trait composition (i.e. dominant traits and functional diversity) of diverse bee communities in contrasting fire-driven systems in two climatic regions: Mediterranean (scrub habitats in Israel) and temperate (chestnut forests in southern Switzerland). 3. In both climatic regions, there were shifts in species diversity and composition related to post-fire age. In the temperate region, functional composition responded markedly to fire; however, in the Mediterranean, the taxonomic response to fire was not matched by functional replacement. 4. These results suggest that greater functional stability to fire in the Mediterranean is achieved by replacement of functionally similar species (i.e. functional redundancy) which dominate under different environmental conditions in the heterogeneous landscapes of the region. In contrast, the greater functional response in the temperate region was attributed to a more rapid post-fire vegetation recovery and shorter time-window when favourable habitat was available relative to the Mediterranean. 5. Bee traits can be used to predict the functional responses of bee communities to environmental changes in habitats of conservation importance in different regions with distinct disturbance regimes. However, predictions cannot be generalized from one climatic region to another where distinct habitat configurations occur.     

Species richness increases the resilience of wetland plant communities in a tropical floodplain

In the last two decades, the relationship between diversity and stability/ecosystem functioning has been widely discussed and has become a central issue in ecology. Here, we assessed the relationship between wetland plant diversity and community resilience after a disturbance. Our study area was located in the Upper Paraná River floodplain (Brazil). An experiment was carried out in situ (18 1 m × 1 m plots with richness varying from 1 to 18 species). In each plot, we recorded the number of species, total per cent vegetation cover and per cent age cover of each species. The above‐ground biomass of wetland plants was removed, simulating a disturbance by animal trampling or an extreme flood. The recovery of vegetation was monitored over 3 months. According to a linear regression, the recovery of wetland plants was positively correlated with diversity. Comparisons with plots containing monocultures of one of the dominant species (Polygonum stelligerum) suggested that this species did not overyield in mixed cultures. Thus, our experiments indicate that the higher resilience in richer plots after a disturbance is mainly due to the fact that species have different resource use requirements (complementarity effect) and not due to the presence of a single, more productive species. Our experiment carried out in a more real condition (in situ) showed that biodiversity is important to wetland functioning and stability, paralleling the results obtained in laboratory and mesocosms experiments. These results also suggest that the loss of plant diversity in our study area could compromise community recovery following strong disturbances.     

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.     

Recovery of forest soil fauna diversity and composition after repeated summer droughts

To examine the resilience of soil animal communities to large-scale disturbances. we studied the recovery of total abundance, diversity and community composition of forest soil mesofauna after a 6-year climatic disturbance. This was done in a pre-established experiment in a Norway spruce Picea abies stand in southern Sweden in which long-term summer droughts had been experimentally imposed and had caused large changes in soil fauna communities. We included both predators (mesostigmatid mites) and fungivores/detritivores (oribatid mites, collembolans) in the study because of the likelihood that they would differ in recovery ability due to differences in their feeding habits, dispersal ability and reproductive strategies. Total abundances of Collembola, Oribatida and Mesostigmata were similar in recovery and control plots after three years, but species richness, the Shannon-Wiener diversity index, and community composition recovered more slowly, particularly among the Oribatida. To only use total abundance of higher taxonomic groups was thus not sufficient when measuring community recovery. There was a tendency for more mobile groups to recover faster than the slow-moving oribatids, indicating the importance of dispersal ability for the resilience of soil communities.     

Macrofaunal Functional Diversity Provides Resilience to Nutrient Enrichment in Coastal Sediments

The degradation of ecosystems is often associated with losses of large organisms and the concomitant losses of the ecological functions they mediate. Conversely, the resilience of ecosystems to stress is strongly influenced by faunal communities and their impacts on processes. Denitrification in coastal sediments is a process that may provide ecosystem resilience to eutrophication by removing excess bioavailable nitrogen. Here, we conducted a large-scale field experiment to test the effect of macrofaunal community composition on denitrification in response to two levels of nutrient enrichment at 28 sites across a biologically heterogeneous sandflat. After 7 weeks of enrichment, we measured denitrification enzyme activity (DEA) along with benthic macrofaunal community composition and environmental variables. We normalised treatment site specific DEA values by those in ambient sediments (DEA_CN) to reveal the underlying response across the heterogeneous landscape. Nutrient enrichment caused reductions in DEA_CN as well as functional changes in the community; these were both more pronounced under the highest level of nutrient loading (on average DEA_CN was reduced by 34%). The degree of suppression of DEA_CN following moderate nitrogen loading was mitigated by a key bioturbating species, but following high nitrogen loading (which reduced the key species density) the abundance and diversity of other nutrient processing species were the most important factors alleviating negative effects. This study provides a prime example of the context-dependent role of biodiversity in maintaining ecosystem functioning, underlining that different elements of biodiversity can become important as stress levels increase. Our results emphasise that management and conservation strategies require a real-world understanding of the community attributes that facilitate nutrient processing and maintain resilience in coastal ecosystems.     

Pyrodiversity promotes interaction complementarity and population resistance

Theory predicts that network characteristics may help anticipate how populations and communities respond to extreme climatic events, but local environmental context may also influence responses to extreme events. For example, altered fire regimes in many ecosystems may significantly affect the context for how species and communities respond to changing climate. In this study, I tested whether the responses of a pollinator community to extreme drought were influenced by the surrounding diversity of fire histories (pyrodiversity) which can influence their interaction networks via changing partner availability. I found that at the community level, pyrodiverse landscapes promote functional complementarity and generalization, but did not consistently enhance functional redundancy or resistance to simulated co‐extinction cascades. Pyrodiversity instead supported flexible behaviors that enable populations to resist perturbations. Specifically, pollinators that can shift partners and network niches are better able to take advantage of the heterogeneity generated by pyrodiversity, thereby buffering pollinator populations against changes in plant abundances. These findings suggest that pyrodiversity is unlikely to improve community‐level resistance to droughts, but instead promotes population resistance and community functionality. This study provides unique evidence that resistance to extreme climatic events depends on both network properties and historical environmental context.     

Effects of Local Tree Diversity on Herbivore Communities Diminish with Increasing Forest Fragmentation on the Landscape Scale

Forest fragmentation and plant diversity have been shown to play a crucial role for herbivorous insects (herbivores, hereafter). In turn, herbivory-induced leaf area loss is known to have direct implications for plant growth and reproduction as well as long-term consequences for ecosystem functioning and forest regeneration. So far, previous studies determined diverging responses of herbivores to forest fragmentation and plant diversity. Those inconsistent results may be owed to complex interactive effects of both co-occurring environmental factors albeit they act on different spatial scales. In this study, we investigated whether forest fragmentation on the landscape scale and tree diversity on the local habitat scale show interactive effects on the herbivore community and leaf area loss in subtropical forests in South Africa. We applied standardized beating samples and a community-based approach to estimate changes in herbivore community composition, herbivore abundance, and the effective number of herbivore species on the tree species-level. We further monitored leaf area loss to link changes in the herbivore community to the associated process of herbivory. Forest fragmentation and tree diversity interactively affected the herbivore community composition, mainly by a species turnover within the family of Curculionidae. Furthermore, herbivore abundance increased and the number of herbivore species decreased with increasing tree diversity in slightly fragmented forests whereas the effects diminished with increasing forest fragmentation. Surprisingly, leaf area loss was neither affected by forest fragmentation or tree diversity, nor by changes in the herbivore community. Our study highlights the need to consider interactive effects of environmental changes across spatial scales in order to draw reliable conclusions for community and interaction patterns. Moreover, forest fragmentation seems to alter the effect of tree diversity on the herbivore community, and thus, has the potential to jeopardize ecosystem functioning and forest regeneration.     

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.