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13 pages, 5 figures, 5 tables, supporting information https://doi.org/10.1002/ece3.9929.-- Data Availability Statement: The data used in this study will be made publicly available at the Dryad open repository (https://datadryad.org/stash), Abiotic environmental conditions can significantly influence the way species interact. In particular, plant–herbivore interactions can be substantially dependent on temperature and nutrients. The overall product of these relationships is critical for the fate and stability of vegetated ecosystems like marine forests. The last few decades have seen a rapid spread of barrens on temperate rocky reefs mainly as a result of overgrazing. The ecological feedbacks that characterize the barren state involve a different set of interactions than those occurring in vegetated habitats. Reversing these trends requires a proper understanding of the novel feedbacks and the conditions under which they operate. Here, we explored the role of a secondary herbivore in reinforcing the stability of barrens formed by sea urchin overgrazing under different nutrient conditions. Combining comparative and experimental studies in two Mediterranean regions characterized by contrasting nutrient conditions, we assessed: (i) if the creation of barren areas enhances limpet abundance, (ii) the size-specific grazing impact by limpets, and (iii) the ability of limpets alone to maintain barrens. Our results show that urchin overgrazing enhanced limpet abundance. The effects of limpet grazing varied with nutrient conditions, being up to five times more intense under oligotrophic conditions. Limpets were able to maintain barrens in the absence of sea urchins only under low-nutrient conditions, enhancing the stability of the depauperate state. Overall, our study suggests a greater vulnerability of subtidal forests in oligotrophic regions of the Mediterranean and demonstrates the importance of environment conditions in regulating feedbacks mediated by plant–herbivore interactions, J Boada acknowledges funding from the Spanish Ministry of Science and Innovation (FJC2018-035566-I) and the European Commission – European Union's Horizon 2020 MSCA – SHIFT2SOLVE-1030591. T Alcoverro acknowledges funding from the Spanish Ministry of Science and Innovation – Project UMBRAL (CTM2017-86695-C3-1R). F. Bulleri acknowledges support from the FutureMARES project, funded from the European Union's Horizon 2020 research and innovation program under grant agreement No. 869300, With the institutional support of the ‘Severo Ochoa Centre of Excellence’ accreditation (CEX2019-000928-S)

1. Theory in ecology and evolution often relies on the analysis of invasion processes, and general approaches exist to understand the early stages of an invasion. However, predicting the long-term transformations of communities following an invasion remains a challenging endeavour. 2. We propose a general analytical method that uses both resident community and invader dynamical features to predict whether an invasion causes large long-term impacts on the invaded community. 3. This approach reveals a direction in which classic invasion analysis, based on initial invasion growth rate, can be extended. Indeed, we explain how the density dependence of invasion growth, if properly defined, synthetically encodes the long-term biotic transformations caused by an invasion, and therefore predicts its ultimate outcome. This approach further clarifies how the density dependence of the invasion growth rate is as much a property of the invading population as it is one of the invaded community. 4. Our theory applies to any stable community model, and directs us towards new questions that may enrich the toolset of invasion analysis, and suggests that indirect interactions and dynamical stability are key determinants of invasion outcomes. Funding Agencies|H2020 European Research Council [666971]; Agence Nationale de la RechercheFrench National Research Agency (ANR)European Commission [ANR-10-LABX-41]; VetenskapsradetSwedish Research Council [VR-2017-05245]; Irish Research CouncilIrish Research Council for Science, Engineering and Technology [IRCLA/2017/186]

© 2021 The Authors., It is common to characterise the species niche using climate and global species distribution maps. This is then used to predict changes in distribution under a warming climate. This approach assumes that climate is a major driver of species distribution and that each species responds individually (sensu Gleason) to climate. However, in many world landscapes, for a given climate, strikingly different vegetation types co-occur: forests and non-forests. These two alternative biome states are maintained by different feedback processes and have radically different species with contrasting shade and disturbance tolerance traits. We propose that to improve predictions of species distribution changes under a novel climate, we need to consider the presence or absence of forest shade, as species are likely to respond individually only within their forest or non-forest biome, and not across biomes. Synthesis. By considering shade as a biotic filter in niche modelling, we are not only improving our predictive capacity, but we are also reconciling the two views of communities: both the individualistic (within biome) and the organismal (across biomes) views of the community concept become relevant and complementary., This research has been performed under the framework of the projects FIROTIC (PGC2018-096569-B-I00, Spanish government).

[Background] In aquatic ecosystems, the health and performance of fish depend greatly on the dynamics of microbial community structure in the background environment. Nonetheless, finding microbes with profound impacts on fish’s performance out of thousands of candidate species remains a major challenge. [Methods] We examined whether time-series analyses of microbial population dynamics could illuminate core components and structure of fish-associated microbiomes in the background (environmental) water. By targeting eel-aquaculture-tank microbiomes as model systems, we reconstructed the population dynamics of the 9605 bacterial and 303 archaeal species/strains across 128 days. [Results] Due to the remarkable increase/decrease of constituent microbial population densities, the taxonomic compositions of the microbiome changed drastically through time. We then found that some specific microbial taxa showed a positive relationship with eels’ activity levels even after excluding confounding effects of environmental parameters (pH and dissolved oxygen level) on population dynamics. In particular, a vitamin-B12-producing bacteria, Cetobacterium somerae, consistently showed strong positive associations with eels’ activity levels across the replicate time series of the five aquaculture tanks analyzed. Network theoretical and metabolic modeling analyses further suggested that the highlighted bacterium and some other closely-associated bacteria formed “core microbiomes” with potentially positive impacts on eels. [Conclusions] Overall, these results suggest that the integration of microbiology, ecological theory, and network science allows us to explore core species and interactions embedded within complex dynamics of fish-associated microbiomes., 魚の健康において鍵となる「コア微生物叢」 --ウナギ養殖水槽内の細菌叢動態--. 京都大学プレスリリース. 2023-03-30.

Active restoration is frequently implemented to restore degraded drylands globally, yet predicting the overall success of these restoration projects remains challenging. Here, we aim to explore if vegetation spatial patterns can be used to monitor ecosystem recovery and anticipate the success of restoration practices. We combined field surveys and high-resolution drone images to analyze how vegetation spatial patterns change after the large-scale straw checkerboards restoration projects in the sand dune systems of the Tengger Desert in northern China. We found that vegetation cover, plant species richness, and diversity increased rapidly, approaching the level of the naturally vegetated sand dunes, after 5 years of restoration. Soil fertility remained low despite the positive vegetation change. Along with the recovery process, we found a larger diversity of patch sizes (i.e. spatial variance) with an increasing proportion of large-size patches. These patterns, combined with a constant positive recovery rate in vegetation cover, are consistent with theoretical predictions used to anticipate transitions to alternative ecosystem states. Although some indicators may be masked by the artificial planting scheme and community succession process during recovery, our results show that vegetation spatial patterns can be used to forecast and monitor the possible recovery of drylands.

Anthropogenic activities are increasingly affecting ecosystems across the globe. Meanwhile, empirical and theoretical evidence suggest that natural systems can exhibit abrupt collapses in response to incremental increases in the stressors, sometimes with dramatic ecological and economic consequences. These catastrophic shifts are faster and larger than expected from the changes in the stressors and happen once a tipping point is crossed. The primary mechanisms that drive ecosystem responses to perturbations lie in their architecture of relationships, i.e. how species interact with each other and with the physical environment and the spatial structure of the environment. Nonetheless, existing theoretical work on catastrophic shifts has so far largely focused on relatively simple systems that have either few species and/or no spatial structure. This work has laid a critical foundation for understanding how abrupt responses to incremental stressors are possible, but it remains difficult to predict (let alone manage) where or when they are most likely to occur in more complex real-world settings. Here, we discuss how scaling up our investigations of catastrophic shifts from simple to more complex—species rich and spatially structured—systems could contribute to expanding our understanding of how nature works and improve our ability to anticipate the effects of global change on ecological systems. This article is part of the theme issue ‘Ecological complexity and the biosphere: the next 30 years’.

Significance Changes in ocean redox chemistry are frequently observed in Earth’s history and have fundamental implications for the evolution of marine life. These transitions are commonly ascribed to large changes in the supply of iron, sulfur, or organic carbon in the deeper ocean. We propose that small variations in carbon input flux can drive nonreversible redox changes of the ocean interior and other anoxic systems, such as marine sediments. Nonlinear interactions in the iron and sulfur cycles create tipping points where regime shifts can occur between alternative stable states that are either iron dominated or sulfide dominated. The recognition that the biogeochemistry of sediments and oceans embeds intrinsic bistability provides a conceptual framework for understanding past and present anoxic marine systems., For most of Earth’s history, the ocean’s interior was pervasively anoxic and showed occasional shifts in ocean redox chemistry between iron-buffered and sulfide-buffered states. These redox transitions are most often explained by large changes in external inputs, such as a strongly altered delivery of iron and sulfate to the ocean, or major shifts in marine productivity. Here, we propose that redox shifts can also arise from small perturbations that are amplified by nonlinear positive feedbacks within the internal iron and sulfur cycling of the ocean. Combining observational evidence with biogeochemical modeling, we show that both sedimentary and aquatic systems display intrinsic iron–sulfur bistability, which is tightly linked to the formation of reduced iron–sulfide minerals. The possibility of tipping points in the redox state of sediments and oceans, which allow large and nonreversible geochemical shifts to arise from relatively small changes in organic carbon input, has important implications for the interpretation of the geological rock record and the causes and consequences of major evolutionary transitions in the history of Earth’s biosphere.

There is growing interest in the application of alternative stable state (ASS) theory to explain major vegetation patterns of the world. Here, we introduce the theory as applied to the puzzle of nonforested (open) biomes growing in climates that are warm and wet enough to support forests (alternative biome states, ABSs). Long thought to be the product of deforestation, diverse lines of evidence indicate that many open ecosystems are ancient. They have also been characterized as ‘early successional’ even where they persist for millennia. ABS is an alternative framework to that of climate determinism and succession for exploring forest/nonforest mosaics. This framework explains not only tropical forest–savanna landscapes, but also other landscape mosaics across the globe.

Humans were considered external drivers in much foundational ecological research. A recognition that humans are embedded in the complex interaction networks we study can provide new insight into our ecological paradigms. Here, we use time-series data spanning three decades to explore the effects of human harvesting on otter–urchin–kelp trophic cascades in southeast Alaska. These effects were inferred from variation in sea urchin and kelp abundance following the post fur trade repatriation of otters and a subsequent localized reduction of otters by human harvest in one location. In an example of a classic trophic cascade, otter repatriation was followed by a 99% reduction in urchin biomass density and a greater than 99% increase in kelp density region wide. Recent spatially concentrated harvesting of otters was associated with a localized 70% decline in otter abundance in one location, with urchins increasing and kelps declining in accordance with the spatial pattern of otter occupancy within that region. While the otter–urchin–kelp trophic cascade has been associated with alternative community states at the regional scale, this research highlights how small-scale variability in otter occupancy, ostensibly due to spatial variability in harvesting or the risk landscape for otters, can result in within-region patchiness in these community states.

Detecting alternative attractors in ecosystem dynamics This repository contains all code needed to reproduce the analyses conducted in the manuscript "Detecting alternative attractors in ecosystem dynamics" (Säterberg & McCann 2021 Communications Biology). Abstract: Dynamical systems theory suggests that ecosystems may exhibit alternative dynamical attractors. Such alternative attractors, as for example equilibria and cycles, have been found in the dynamics of experimental systems. Yet, for natural systems, where multiple biotic and abiotic factors simultaneously affect population dynamics, it is more challenging to distinguish alternative dynamical behaviors. Although recent research exemplifies that some natural systems can exhibit alternative states, a robust methodology for testing whether these constitute distinct dynamical attractors is currently lacking. Here, using attractor reconstruction techniques we develop such a test. Applications of the methodology to simulated, experimental and natural time series data, reveal that alternative dynamical behaviors are hard to distinguish if population dynamics are governed by purely stochastic processes. However, if population dynamics are brought about also by mechanisms internal to the system, alternative attractors can readily be detected. Since many natural populations display evidence of such internally-driven dynamics, our approach offers a method for empirically testing whether ecosystems exhibit alternative dynamical attractors., We thank Gregor Fussmann for providing experimental data that was used to test the methodology in this publication.

Globally, peatlands have been affected by drainage and peat extraction, with adverse effects on their functioning and services. To restore peat-forming vegetation, drained bogs are being rewetted on a large scale. Although this practice results in higher groundwater levels, unfortunately it often creates deep lakes in parts where peat was extracted to greater depths than the surroundings. Revegetation of these deeper waters by peat mosses appears to be challenging due to strong abiotic feedbacks that keep these systems in an undesired bare state. In this study, we theoretically explore if a floating peat mat and an open human-made bog lake can be considered two alternative stable states using a simple model, and experimentally test in the field whether stable states are present, and whether a state shift can be accomplished using floating biodegradable structures that mimic buoyant peat. We transplanted two peat moss species into these structures (pioneer sp. Sphagnum cuspidatum and later-successional sp. S. palustre) with and without additional organic substrate. Our model suggests that these open human-made bog lakes and floating peat mats can indeed be regarded as alternative stable states. Natural recovery by spontaneous peat moss growth, i.e., a state shift from open water to floating mats, is only possible when the water table is sufficiently shallow to avoid light limitation (S. cuspidatum and vascular plants. Organic substrate addition particularly facilitated vascular plant growth, which correlated to higher moss height. The structures remained too wet for the late-successional species S. palustre. We conclude that open water and floating peat mats in human-made bog lakes can be considered two alternative stable states, and that temporary floating establishment structures can induce a state shift from the open water state to peat-forming vegetation state. These findings imply that for successful restoration, there is a clear water depth threshold to enable peat moss growth and there is no need for addition of large amounts of donor-peat substrate. Correct species selection for restoration is crucial for success.

Ecosystems worldwide depend on habitat‐forming foundation species that often facilitate themselves with increasing density and patch size, while also engaging in facultative mutualisms. Anthropogenic global change (e.g., climate change, eutrophication, overharvest, land‐use change), however, is causing rapid declines of foundation species‐structured ecosystems, often typified by sudden collapse. Although disruption of obligate mutualisms involving foundation species is known to precipitate collapse (e.g., coral bleaching), how facultative mutualisms (i.e., context‐dependent, nonbinding reciprocal interactions) affect ecosystem resilience is uncertain. Here, we synthesize recent advancements and combine these with model analyses supported by real‐world examples, to propose that facultative mutualisms may pose a double‐edged sword for foundation species. We suggest that by amplifying self‐facilitative feedbacks by foundation species, facultative mutualisms can increase foundation species’ resistance to stress from anthropogenic impact. Simultaneously, however, mutualism dependency can generate or exacerbate bistability, implying a potential for sudden collapse when the mutualism's buffering capacity is exceeded, while recovery requires conditions to improve beyond the initial collapse point (hysteresis). Thus, our work emphasizes the importance of acknowledging facultative mutualisms for conservation and restoration of foundation species‐structured ecosystems, but highlights the potential risk of relying on mutualisms in the face of global change. We argue that significant caveats remain regarding the determination of these feedbacks, and suggest empirical manipulation across stress gradients as a way forward to identify related nonlinear responses., Ecosystems worldwide depend on habitat‐forming foundation species that engage in facultative mutualisms. Global change, however, is causing rapid declines of foundation species‐structured ecosystems, often typified by sudden collapse. Here, we synthesize recent advancements and combine this with model analyses to propose that facultative mutualisms can increase foundation species resistance to global change stressors, but simultaneously increase the risk of sudden collapse by generating or exacerbating bistability.

© 2021 The Authors. It is common to characterise the species niche using climate and global species distribution maps. This is then used to predict changes in distribution under a warming climate. This approach assumes that climate is a major driver of species distribution and that each species responds individually (sensu Gleason) to climate. However, in many world landscapes, for a given climate, strikingly different vegetation types co-occur: forests and non-forests. These two alternative biome states are maintained by different feedback processes and have radically different species with contrasting shade and disturbance tolerance traits. We propose that to improve predictions of species distribution changes under a novel climate, we need to consider the presence or absence of forest shade, as species are likely to respond individually only within their forest or non-forest biome, and not across biomes. Synthesis. By considering shade as a biotic filter in niche modelling, we are not only improving our predictive capacity, but we are also reconciling the two views of communities: both the individualistic (within biome) and the organismal (across biomes) views of the community concept become relevant and complementary. This research has been performed under the framework of the projects FIROTIC (PGC2018-096569-B-I00, Spanish government).

The expansion of rainforest pioneer trees into long-unburnt open forests is a global phenomenon and widely reported in high rainfall regions of Australia. However, impacts on Australian open forest biodiversity have been little studied beyond the tropics. In this thesis, I asked whether an invading midstorey of rainforest pioneers had adverse impacts on communities of open forest flora and fauna, and on ecosystem function, by examining heathy Eucalypt forest with contrasting fire frequencies and levels of rainforest-invasion in the Bundjalung Wilderness Area of northern NSW., Where rainforest pioneers dominated the midstorey, understorey plant species richness was halved, and ground cover and density of open forest specialists were reduced by ~90% compared to the uninvaded state. Rainforest-invaded open forest also had lower insectivorous bat activity (63% lower) and species richness (35% lower). Forest flammability was also sharply reduced by rainforest pioneers through near-elimination of dense grassy (graminoid) and heathy understorey fuels in exchange for sparse rainforest saplings with less-flammable foliage and crown structure., These findings suggest that rainforest pioneers are ecosystem engineers that modify open forest fuel properties, initiating a positive fire-suppression feedback that facilitates their persistence and the continued transition to a fire-resistant closed forest., I also examined fire-resilience traits among 228 taxa of woody rainforest plants burnt in the extensive 2019-20 bushfires of eastern Australia to determine the reversibility of rainforest-invasion when fire ultimately returns to long-unburnt open forest. Most taxa survived fire by resprouting and/or fire-cued seedling recruitment, and resprouting rates were high across all size classes, including in small rainforest plants (���1 cm DBH, ���1m tall)., These findings suggest that rainforest plants which invade long-unburnt open forests may quickly become resistant to removal by future fires, allowing rapid re-establishment of midstorey cover from established root stocks., The observed declines in open forest flora, fauna and ecosystem function occurred well before recommended maximum fire-intervals for dry open forest, casting doubt on the suitability of existing guidelines in high rainfall regions. To conserve open forest biodiversity, this research suggests that fires should be of sufficient frequency to prevent a dense rainforest midstorey from eliminating open forest flora, fauna and the understorey fuels that underpin open forest function.

Global forests are experiencing widespread climate-induced mortality. Predicting this phenomenon has proven difficult, despite recent advances in understanding physiological mechanisms of mortality in individual trees along with environmental drivers of mortality at broad scales. With heat and drought as primary climatic drivers, and convergence on hydraulic failure as a primary physiological mechanism, new models are needed to improve our predictions of Earth’s forests under future climate conditions. While much of ecology focuses on equilibrium states, transitions from one stable state to another are often described with alternative stable state theory (ASST), where systems can settle to more than one stable condition. Recent studies have identified threshold responses of hydraulic failure during tree mortality, indicating that alternative stable states may be present. Here, I demonstrate that the xylem of trees has characteristics indicative of alternative stable states. Through empirical evidence, I identify a catastrophic shift during hydraulic failure which prevents trees from returning to pre-droughted physiological states after environmental stressors (e.g., drought, heat) are relieved. Thus, the legacy of climate-induced hydraulic failure likely contributes to reduced resilience of forests under future climate. I discuss the implications and future directions for including ASST in models of tree mortality.

Significance Intrinsic institutional or ecological differences are often invoked to explain resource conservation success or failure. However, alternative conservation outcomes instead may be caused by path-dependent processes, where historical contingencies trap similar institutions in dramatically different, but predictable, states. We model social-ecological processes in cooperatively managed natural resources and show that institutional path dependence can create alternative stable states of conservation or overharvesting. We find that the model significantly explains a large dataset of well-studied marine fisheries. Highly productive and costly resources are, unexpectedly, most likely to exhibit the alternative stable states of strong depletion or conservation. Path dependence presents challenges and opportunities, including the possibility that short but intensive harvest reduction efforts can generate self-perpetuating conservation outcomes., Understanding why some renewable resources are overharvested while others are conserved remains an important challenge. Most explanations focus on institutional or ecological differences among resources. Here, we provide theoretical and empirical evidence that conservation and overharvest can be alternative stable states within the same exclusive-resource management system because of path-dependent processes, including slow institutional adaptation. Surprisingly, this theory predicts that the alternative states of strong conservation or overharvest are most likely for resources that were previously thought to be easily conserved under optimal management or even open access. Quantitative analyses of harvest rates from 217 intensely managed fisheries supports the predictions. Fisheries’ harvest rates also showed transient dynamics characteristic of path dependence, as well as convergence to the alternative stable state after unexpected transitions. This statistical evidence for path dependence differs from previous empirical support that was based largely on case studies, experiments, and distributional analyses. Alternative stable states in conservation appear likely outcomes for many cooperatively managed renewable resources, which implies that achieving conservation outcomes hinges on harnessing existing policy tools to navigate transitions.

Fires and herbivores shape tropical vegetation structure, but their effects on the stability of tree cover in different climates remains elusive. Here we integrate empirical and theoretical approaches to determine the effects of climate on fire- and herbivore-driven forest-savanna shifts. We analyzed time series of remotely sensed tree cover and fire observations with estimates of herbivore pressure across the tropics to quantify the fire-tree cover and herbivore-tree cover feedbacks along climatic gradients. From these empirical results we developed a spatially explicit, stochastic fire-vegetation model that accounts for herbivore pressure. We find emergent alternative stable states in tree cover with hysteresis across rainfall conditions. Whereas the herbivore-tree cover feedback can maintain low tree cover below 1100 mm mean annual rainfall, the fire-tree cover feedback can maintain low tree cover at higher rainfall levels. Interestingly, the rainfall range where fire-driven alternative vegetation states can be found depends strongly on rainfall variability. Both higher seasonal and inter-annual variability in rainfall increase fire frequency, but only seasonality expands the distribution of fire-maintained savannas into wetter climates. The strength of the fire-tree cover feedback depends on the spatial configuration of tree cover: landscapes with clustered low tree cover areas are more susceptible to cross a tipping point of fire-driven forest loss than landscapes with scattered deforested patches. Our study shows how feedbacks involving fire, herbivores and the spatial structure of tree cover explain the resilience of tree cover across climates. This article is protected by copyright. All rights reserved.

International audience; Symbiotic nitrogen (N)‐fixing plants are abundant during primary succession, as typical bedrocks lack available N. In turn, fixed N accumulates in soils through biomass turnover and recycling, favouring more nitrophilous organisms. Yet, it is unclear how this facilitation mechanism interacts with competition for other limiting nutrients such as phosphorus (P) and how this affects succession. Here, we introduce a resource‐explicit, community assembly model of N‐fixing species and analyze successional trajectories along resource availability gradients using contemporary niche theory. We show that facilitation‐driven succession occurs under low N and high enough P availabilities, and is characterised by autogenic ecosystem development and relatively ordered trajectories. We show that late facilitation‐driven succession is sensitive to catastrophic shifts, highlighting the need to invoke other mechanisms to explain ecosystem stability near the climax. Put together with competition‐driven succession, these results lead to an enriched version of Tilman's resource‐ratio theory of succession.

Aim: Although much tropical ecology generally focuses on trees, grasses are fundamental for characterizing the extensive tropical grassy biomes (TGBs) and, together with the tree functional types, for determining the contrasting functional patterns of TGBs and tropical forests (TFs). To study the factors that determine African biome distribution and the transitions between them, we performed the first continental analysis to include grass and tree functional types. Location: Sub-Saharan Africa. Time period: 2000-2010. Major taxa studied: Savanna and forest trees and C4 grasses. Methods: We combined remote-sensing data with a land cover map, using tree functional types to identify TGBs and TFs. We analysed the relationships of grass and tree cover with fire interval, rainfall annual average and seasonality. Results: In TGBs experiencing < 630 mm annual rainfall, grass growth was water limited. Grass cover and fire recurrence were strongly and directly related over the entire subcontinent. Some TGBs and TFs with annual rainfall > 1,200 mm had the same rainfall seasonality but displayed strongly different fire regimes. Main conclusions: Water limitation to grass growth was fundamental in the driest TGBs, acting alongside the well-known limitation to tree growth. Marked differences in fire regimes across all biomes indicated that fire was especially relevant for maintaining mesic and humid TGBs. At high rainfall, our results support the hypothesis of TGBs and TFs being alternative stable states maintained by a vegetation-fire feedback for similar climatic conditions.

© 2017 by the Ecological Society of America. A major goal of ecosystem-based fisheries management is to prevent fishery-induced shifts in community states. This requires an understanding of ecological resilience: the ability of an ecosystem to return to the same state following a perturbation, which can strongly depend on species interactions across trophic levels. We use a structured model of a temperate rocky reef to explore how multi-trophic level fisheries impact ecological resilience. Increasing fishing mortality of prey (urchins) has a minor effect on equilibrium biomass of kelp, urchins, and spiny lobster predators, but increases resilience by reducing the range of predator harvest rates at which alternative stable states are possible. Size-structured predation on urchins acts as the feedback maintaining each state. Our results demonstrate that the resilience of ecosystems strongly depends on the interactive effects of predator and prey harvest in multi-trophic level fisheries, which are common in marine ecosystems but are unaccounted for by traditional management.

There are strong opposing views among conservationists about whether we have to intervene to safeguard our natural heritage or not. In the Western European tradition, human intervention has been dominating, whereas, elsewhere, rewilding aimed at restoring self-regulating ecosystems has often been preferred. However, cultural rather than ecological differences are at the root of these opposing paradigms, leading to management strategies that are not always optimal for biodiversity conservation. In the present article, we propose a framework based on the relationship between ecosystem dynamics and the human footprint, including land-use legacies, to guide the mixture of rewilding and intervention practices in order to ensure a biodiverse future. We argue that these paradigms are not conflicting but complementary and advocate for rewilding where possible, human intervention where needed.

It is hypothesised that ecological restoration in grasslands can induce an alternative stable state shift in vegetation. The change in vegetation influences insect community assemblages and allows for greater functional redundancy in pollination and refuge for native insect species. 2. Insect community assemblages at eight coastal California grassland sites were evaluated. Half of these sites had undergone restoration through active revegetation of native grassland flora and half were non-restored. Insects were collected from Lupinus bicolor (Fabaceae) within 2 × 2-m2 plots in spring 2017. Lupinus bicolor is a common native species that is used in California restoration projects, and home and state landscaping projects. 3. Ordination demonstrated that insect community assemblages were different between restored and non-restored sites. These differences were seen in insect functional groups as well as taxa-specific differences and were found to be driven by environmental characteristics such as non-native forb cover. 4. Functional redundancy of herbivores decreased at restored sites, while pollinators became more redundant compared with non-restored sites. The assemblages of the common species found at restoration sites contained more native insects than those found at non-restored sites, including species such as Bombus vosnesenskii. 5. Local grassland restoration has the potential to induce an alternative stable state change and affect insect community assemblages. Additionally, it was found that grassland restoration can be a potential conservation tool to provide refugia for bumblebees (Bombus), but additional studies are required to fully understand its broader applicability.

Glacier melting sediment inputs affect coastal ecosystems on the Antarctic Peninsula. In Potter Cove (South Shetland Islands, Antarctica), the shift from an “ascidian dominated” to a “mixed” assemblage has been linked to sedimentation. However, in recently described newly ice-free areas ascidians became dominant in spite of total suspended particulate matter (TSPM) concentrations, which are the highest measured in Potter Cove. Here, we compared the gut content and energy reserve of three ascidian species at three stations under different TSPM regimes. All analysed species had a higher gut content with lower %OM at these newly areas. A theoretical relationship between the scope for growth for the targeted ascidians and TSPM explained assemblages’ recorded change but failed to explain current ascidians distribution. The results may indicate the existence of a TSPM threshold that allows the spatial coexistence of alternative stable states at benthic Potter Cove system. Fil: Torre, Luciana. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto de Diversidad y Ecología Animal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto de Diversidad y Ecología Animal; Argentina Fil: Alurralde, Roque Gastón. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto de Diversidad y Ecología Animal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto de Diversidad y Ecología Animal; Argentina Fil: Lagger, Cristian Fabian. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto de Diversidad y Ecología Animal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto de Diversidad y Ecología Animal; Argentina Fil: Abele, D.. Helmholtz Centre for Polar and Marine Research; Alemania Fil: Schloss, Irene Ruth. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentina Fil: Sahade, Ricardo Jose. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto de Diversidad y Ecología Animal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto de Diversidad y Ecología Animal; Argentina

Although canopy height has long been a focus of interest in ecology, it has remained difficult to study at large spatial scales. Recently, satellite-borne LiDAR equipment produced the first systematic high resolution maps of vegetation height worldwide. Here we show that this new resource reveals three marked modes in tropical canopy height ~40, ~12, and ~2 m corresponding to forest, savanna, and treeless landscapes. The distribution of these modes is consistent with the often hypothesized forest-savanna bistability and suggests that both states can be stable in areas with a mean annual precipitation between ~1,500 and ~2,000 mm. Although the canopy height states correspond largely to the much discussed tree cover states, there are differences, too. For instance, there are places with savanna-like sparse tree cover that have a forest-like high canopy, suggesting that rather than true savanna, those are thinned relicts of forest. This illustrates how complementary sets of remotely sensed indicators may provide increasingly sophisticated ways to study ecological phenomena at a global scale.

© 2018 by the Ecological Society of America Woody plant encroachment into grasslands is a major land cover change taking place in many regions of the world, including arctic, alpine and desert ecosystems. This change in plant dominance is also affecting coastal ecosystems, including barrier islands, which are known for being vulnerable to the effects of climate change. In the last century, the woody plant species Morella cerifera L. (Myricaceae), has encroached into grass covered swales in many of the barrier islands of Virginia along the Atlantic seaboard. The abrupt shift to shrub cover in these islands could result from positive feedbacks with the physical environment, though the underlying mechanisms remain poorly understood. We use a combination of experimental and modeling approaches to investigate the role of climate warming and the ability of M. cerifera to mitigate its microclimate thereby leading to the emergence of alternative stable states in barrier island vegetation. Nighttime air temperatures were significantly higher in myrtle shrublands than grasslands, particularly in the winter season. The difference in the mean of the 5% and 10% lowest minimum temperatures between shrubland and grassland calculated from two independent datasets ranged from 1.3 to 2.4°C. The model results clearly show that a small increase in near-surface temperature can induce a non-linear shift in ecosystem state from a stable state with no shrubs to an alternative stable state dominated by M. cerifera. This modeling framework improves our understanding and prediction of barrier island vegetation stability and resilience under climate change, and highlights the existence of important nonlinearities and hystereses that limit the reversibility of this ongoing shift in vegetation dominance.

The spatial dispersal of individuals plays an important role in the dynamics of populations, and is central to metapopulation theory. Dispersal provides connections within metapopulations, promoting demographic and evolutionary rescue, but may also introduce maladapted individuals, potentially lowering the fitness of recipient populations through introgression of heritable traits. To explore this dual nature of dispersal, we modify a well-established eco-evolutionary model of two locally adapted populations and their associated mean trait values, to examine recruiting salmon populations that are connected by density-dependent dispersal, consistent with collective migratory behaviour that promotes navigation. When the strength of collective behaviour is weak such that straying is effectively constant, we show that a low level of straying is associated with the highest gains in metapopulation robustness and that high straying serves to erode robustness. Moreover, we find that as the strength of collective behaviour increases, metapopulation robustness is enhanced, but this relationship depends on the rate at which individuals stray. Specifically, strong collective behaviour increases the presence of hidden low-density basins of attraction, which may serve to trap disturbed populations, and this is exacerbated by increased habitat heterogeneity. Taken as a whole, our findings suggest that density-dependent straying and collective migratory behaviour may help metapopulations, such as in salmon, thrive in dynamic landscapes. Given the pervasive eco-evolutionary impacts of dispersal on metapopulations, these findings have important ramifications for the conservation of salmon metapopulations facing both natural and anthropogenic contemporary disturbances.This article is part of the theme issue 'Collective movement ecology'.

© 2015 by the Ecological Society of America. Extreme events, which have profound ecological consequences, are changing in both frequency and magnitude with climate change. Because extreme temperatures induce coral bleaching, we can explore the relative impacts of changes in frequency and magnitude of high temperature events on coral reefs. Here, we combined climate projections and a dynamic population model to determine how changing bleaching regimes influence coral persistence. We additionally explored how coral traits and competition with macroalgae mediate changes in bleaching regimes. Our results predict that severe bleaching events reduce coral persistence more than frequent bleaching. Corals with low adult mortality and high growth rates are successful when bleaching is mild, but bleaching resistance is necessary to persist when bleaching is severe, regardless of frequency. The existence of macroalgae-dominated stable states reduces coral persistence and changes the relative importance of coral traits. Building on previous studies, our results predict that management efforts may need to prioritize protection of "weaker" corals with high adult mortality when bleaching is mild, and protection of "stronger" corals with high bleaching resistance when bleaching is severe. In summary, future reef projections and conservation targets depend on both local bleaching regimes and biodiversity.

Ecosystems and biological systems are known to be inherently complex and to exhibit nonlinear dynamics. Diseases such as microbiome dysregulation or depression can be seen as complex systems as well and were shown to exhibit patterns of nonlinearity in their response to perturbations. These nonlinearities can be revealed by a sudden shift in system states, for instance from health to disease. The identification and characterization of early warning signals which could predict upcoming critical transitions is of primordial interest as prevention of disease onset is a major aim in health care. In this review, we focus on recent evidence for critical transitions in diseases and discuss the potential of such studies for therapeutic applications.

Intraguild predation interactions make fish communities prone to exhibit alternative stable states with either piscivore or prey fish dominance. In the Baltic Sea, local declines of coastal piscivores like perch (Perca fluviatilis) have been observed to coincide with high densities of sticklebacks (Gasterosteus aculeatus). Mechanisms behind this shift between piscivore and stickleback dominance were studied both experimentally and in field. Results showed that predation by sticklebacks has a strong negative effect on perch larvae survival, but this effect rapidly decreases with increasing perch size, likely due to gape limitations and digestion constraints in sticklebacks. Large spatial and temporal variations in patterns of stickleback migration into perch spawning sites were observed. Whether or not high density of sticklebacks will cause declines in coastal piscivore populations is suggested to depend on the availability of spawning sites in which sticklebacks do not migrate into or arrive late in the reproduction season of coastal piscivores. Electronic supplementary material The online version of this article (doi:10.1007/s13280-015-0665-5) contains supplementary material, which is available to authorized users.

© 2017 by the Ecological Society of America Regime shifts are difficult-to-reverse transitions that occur when an ecosystem reorganizes around a new set of self-reinforcing feedbacks. Regime shifts are predicted to occur when the intensity of some exogenous driver variable, such as temperature, annual harvest rate, or nutrient addition rate, gradually approaches and crosses a threshold value, initiating a transition to an alternative state. However, many driver variables now change rapidly as presses or pulses, not gradually, requiring new conceptual frameworks for understanding and predicting regime shifts. We argue that identifying and controlling regime shifts in response to presses and pulses will require a greater focus on the duration, not just the intensity, of changes in driver variables. In ecosystems with slower dynamics, transitions to an alternative state can take years to decades and as a result, a driver press with an intensity capable of resulting in a regime shift over long time spans may fail to cause a regime shift when applied for shorter durations. We illustrate these ideas using simulations of local-scale alternative stable state models and preliminary evidence from long-term grazing and eutrophication experiments. The simulations also suggest that small changes in the duration of driver presses or pulses can determine whether an ecosystem recovers to its original state. These insights may extend to larger scales. In spatially extended simulations that included patchiness, spatial heterogeneity, and spatial connectivity, all patches recovered to their original state after shorter presses. However, once press duration exceeded a threshold, growing proportions of the landscape shifted to an alternative state as press duration increased. We observed similar patchy transitions in a catchment-scale experiment that reinstated frequent fires approximately halfway through a regime shift from grassland to shrubland, initiated by fire suppression. In both the local- and larger-scale models, the threshold duration needed to elicit regime shifts decreased as press intensity increased or when factors counteracting regime shifts weakened. These multiple lines of evidence suggest that conceptualizing regime shifts as an interactive function of the intensity and duration of driver changes will increase understanding of the varying effects of driver presses, pulses, and cycles on ecosystem dynamics.

Este artículo contiene 9 páginas, 5 figuras., Predicting where state-changing thresholds lie can be inherently complex in ecosystems characterized by nonlinear dynamics. Unpacking the mechanisms underlying these transitions can help considerably reduce this unpredictability. We used empirical observations, field and laboratory experiments, and mathematical models to examine how differences in nutrient regimes mediate the capacity of macrophyte communities to sustain sea urchin grazing. In relatively nutrient-rich conditions, macrophyte systems were more resilient to grazing, shifting to barrens beyond 1 800 g m22 (urchin biomass), more than twice the threshold of nutrient-poor conditions. The mechanisms driving these differences are linked to how nutrients mediate urchin foraging and algal growth: controlled experiments showed that low-nutrient regimes trigger compensatory feeding and reduce plant growth, mechanisms supported by our consumer–resource model. These mechanisms act together to halve macrophyte community resilience. Our study demonstrates that by mediating the underlying drivers, inherent conditions can strongly influence the buffer capacity of nonlinear systems., The Spanish Ministry of Science and Innovation funded this research (projects CMT2010-22273-C02-01-02 and CMT2013- 48027-C03-R) and supported J.B. (scholarship BES-2011-043630) and D.A. (Ramon y Cajal fellowship). The Spanish National Research Council supported R.A.’s visitorship (CSIC-201330E062).

When a system has more than one stable state, how can the stability of these states be compared? This deceptively simple question has important consequences for ecosystems, because systems with alternative stable states can undergo dramatic regime shifts. The probability, frequency, duration, and dynamics of these shifts will all depend on the relative stability of the stable states. Unfortunately, the concept of “stability” in ecology has suffered from substantial confusion and this is particularly problematic for systems where stochastic perturbations can cause shifts between coexisting alternative stable states. A useful way to visualize stable states in stochastic systems is with a ball‐in‐cup‐diagram, in which the state of the system is represented as the position of a ball rolling on a surface, and the random perturbations can push the ball from one basin of attraction to another. The surface is determined by a potential function, which provides a natural stability metric. Systems amenable to this representation, called gradient systems, are quite rare, however. As a result, the potential function is not widely used and other approaches based on linear stability analysis have become standard. Linear stability analysis is designed for local analysis of deterministic systems and, as we show, can produce a highly misleading picture of how the system will behave under continual, stochastic perturbations. In this paper, we show how the potential function can be generalized so that it can be applied broadly, employing a concept from stochastic analysis called the quasi‐potential. Using three classic ecological models, we demonstrate that the quasi‐potential provides a useful way to quantify stability in stochastic systems. We show that the quasi‐potential framework helps clarify long‐standing confusion about stability in stochastic ecological systems, and we argue that ecologists should adopt it as a practical tool for analyzing these systems.

It has been suggested that tropical forest and savanna could represent alternative stable states, implying critical transitions at tipping points in response to altered climate or other drivers. So far, evidence for this idea has remained elusive, and integrated climate models assume smooth vegetation responses. We analyzed data on the distribution of tree cover in Africa, Australia, and South America to reveal strong evidence for the existence of three distinct attractors: forest, savanna, and a treeless state. Empirical reconstruction of the basins of attraction indicates that the resilience of the states varies in a universal way with precipitation. These results allow the identification of regions where forest or savanna may most easily tip into an alternative state, and they pave the way to a new generation of coupled climate models.

The population and community level consequences of positive interactions between plants remain poorly explored. In this study we incorporate positive resource-mediated interactions in classic resource competition theory and investigate the main consequences for plant population dynamics and species coexistence. We focus on plant communities for which water infiltration rates exhibit positive dependency on plant biomass and where plant responses can be improved by shading, particularly under water limiting conditions. We show that the effects of these two resource-mediated positive interactions are similar and additive. We predict that positive interactions shift the transition points between different species compositions along environmental gradients and that realized niche widths will expand or shrink. Furthermore, continuous transitions between different community compositions can become discontinuous and bistability or tristability can occur. Moreover, increased infiltration rates may give rise to a new potential coexistence mechanism that we call controlled facilitation. © 2009 Elsevier Inc. All rights reserved.

Thorough understanding of the potential for threshold dynamics and catastrophic shifts to occur in natural systems is of great importance for ecosystem conservation and restoration. However, verifying the presence of alternative stable states, one of the theoretical explanations for sudden shifts in natural systems, has proven to be a major challenge. We examine processes on local and landscape scales in salt-marsh pioneer zones, to assess the presence of alternative stable states in this system. To that end, we investigated the presence of typical characteristics of alternative stable states: bimodality and threshold dynamics. We also studied whether vegetation patches remained stable over long time periods. Analysis of false-color aerial photographs revealed clear bimodality in plant biomass distribution. By transplanting Spartina anglica plants of three different biomass classes on three geographically different marshes, we showed that a biomass threshold limits the establishment of Spartina patches, potentially explaining their patchy distribution. The presence of bimodality and biomass thresholds points to the presence of alternative stable states and the potential for sudden shifts, at small, within-patch scales and on short time scales. However, overlay analysis of aerial photographs from a salt marsh in The Netherlands, covering a time span of 22 years, revealed that there was little long-term stability of patches, as vegetation cover in this area is slowly increasing. Our results suggest that the concept of alternative stable states is applicable to the salt-marsh pioneer vegetation on small spatio-temporal scales. However, the concept does not apply to long-term dynamics of decades or centuries of heterogeneous salt-marsh pioneer zones, as landscape-scale processes may determine the large-scale dynamics of salt marshes. Hence, our results provide the interesting perspective that threshold dynamics may occur in systems with, on the long term, only a single stable state.

The goals of ecosystem-based management (EBM) include protecting ecological resilience, the magnitude of a perturbation that a community can withstand and remain in a given state. As a tool to achieve this goal, no-take marine reserves may enhance resilience by protecting source populations or reduce it by concentrating fishing in harvested areas. Here, we test whether spatial management with marine reserves can increase ecological resilience compared to non-spatial (conventional) management using a dynamic model of a simplified fish community with structured predation and competition that causes alternative stable states. Relative to non-spatial management, reserves increase the resilience of the desired (predator-dominated) equilibrium state in both stochastic and deterministic environments, especially under intensive fishing. As a result, spatial management also increases the feasibility of restoring degraded (competitor-dominated) systems, particularly if combined with culling of competitors or stock enhancement of adult predators.

It has been argued that waterfowl and fish may threaten growth of submerged macrophytes, especially in spring during the early growth phase when plant biomass is low. A small reduction of biomass at that time might delay growth or decrease subsequent productivity. We investigated the impact of waterfowl and large fish on the spring growth of fennel pondweed (Potamogeton pectinatus L.) by employing an exclosure experiment in the macrophyte-dominated clear-water Lake Mogan, Turkey. Birds and large fish were excluded from eight plots and both in situ vegetation and macrophytes kept in pots were compared to eight open plots. Also, to investigate the effect of periphyton on plant growth it was removed from half of the pot plants. Exclusion of waterfowl and fish may decrease predation on macroinvertebrates, which in turn may affect periphyton, and macrophyte growth, why macroinvertebrates also were sampled. Waterfowl density was high (15-70 ind. of coot, Fulica atra L. ha-1), abundance of submerged plants was also high with a surface coverage of 70-80%, and benthivorous fish were present, mainly tench, (Tinca tinca L.) and carp, (Cyprinus carpio L.). Exclusion of waterfowl and large fish did not significantly affect the spring growth of pondweed; neither plants growing in situ nor kept in pots. Removal of periphyton from the plants in the pots did not favour growth. The density of macroinvertebrates was not affected by the exclusion of waterfowl and large fish, but it was positively related to aboveground biomass of fennel pondweed. We suggest that even if waterfowl and large fish are in high densities, their effect on fennel pondweed spring growth in lakes with abundant submerged vegetation, such as Lake Mogan, is low. © Springer 2005. The Research Fund of Middle East Technical University, Ankara, financed this study. The Swedish Institute financially supported HS. Suh-endan Karauz was greatly appreciated for proving the coot data. We thank Fatma Saldiran for editorial assistance, Murat Killi, Levent Burnak and Kerem Ali Boyla for helping in the field and Anders Hargeby and Ola Marklund for valuable comments on an earlier version of this manucript.

▪ Abstract We review the evidence of regime shifts in terrestrial and aquatic environments in relation to resilience of complex adaptive ecosystems and the functional roles of biological diversity in this context. The evidence reveals that the likelihood of regime shifts may increase when humans reduce resilience by such actions as removing response diversity, removing whole functional groups of species, or removing whole trophic levels; impacting on ecosystems via emissions of waste and pollutants and climate change; and altering the magnitude, frequency, and duration of disturbance regimes. The combined and often synergistic effects of those pressures can make ecosystems more vulnerable to changes that previously could be absorbed. As a consequence, ecosystems may suddenly shift from desired to less desired states in their capacity to generate ecosystem services. Active adaptive management and governance of resilience will be required to sustain desired ecosystem states and transform degraded ecosystems into fundamentally new and more desirable configurations.

In tropical landscapes, vegetation patches with contrasting tree densities are distributed as mosaics. However, the locations of patches and densities of trees within them cannot be predicted by climate models alone. It has been proposed that plant-fire feedbacks drive functional thresholds at a landscape scale, thereby maintaining open (savanna) and closed (forest) communities as two distinct stable states. However, there is little rigorous field evidence for this threshold model. Here we aim to provide support for such a model from a field perspective and to analyze the functional and phylogenetic consequences of fire in a Brazilian savanna landscape (Cerrado). We hypothesize that, in tropical landscapes, savanna and forest are two stable states maintained by plant-fire feedbacks. If so, their functional and diversity attributes should change abruptly along a community closure gradient. We set 98 plots along a gradient from open savanna to closed forest in the Brazilian Cerrado and tested for a threshold pattern in nine functional traits, five soil features, and seven diversity indicators. We then tested whether the threshold pattern was associated with different fire regimes. Most community attributes presented a threshold pattern on the savanna-forest transition with coinciding breakpoints. The thresholds separated two community states: (1) open environments with low-diversity communities growing in poor soils and dominated by plants that are highly resistant to high-intensity fires; and (2) closed environments with highly diverse plant communities growing in more fertile soils and dominated by shade-tolerant species that efficiently prevent light from reaching the understory. In addition, each state was associated with contrasting fire regimes. Our results are consistent with the hypothesis that forests and savannas are two coexisting stable states with contrasting patterns of function and diversity that are regulated by fire- plant feedbacks; our results also shed light on the mechanism driving each state. Overall, our results support the idea that fire plays an important role in regulating the distribution of savanna and forest biomes in tropical landscapes. © 2013 by the Ecological Society of America., We are grateful to Fundaçao de Amparo à Pesquisa do Estado de Sao Paulo (FAPESP; process: 2010/01835-0),Coordenaçao de Aperfeiçoamento de Pessoal de Nı́vel Superior (CAPES; process: 1019-11-2), Conselho Nacional de Desenvolvimento Cientı́fico e Tecnológico (CNPq), and the Spanish Government (VIRRA and TREVOL projects, CGL2009- 12048/BOS, CGL2012-39938-C02-01) for financial support and the scholarships granted to the authors.

A pronounced, widespread and persistent regime shift among marine ecosystems is observable on temperate rocky reefs as a result of sea urchin overgrazing. Here, we empirically define regime-shift dynamics for this grazing system which transitions between productive macroalgal beds and impoverished urchin barrens. Catastrophic in nature, urchin overgrazing in a well-studied Australian system demonstrates a discontinuous regime shift, which is of particular management concern as recovery of desirable macroalgal beds requires reducing grazers to well below the initial threshold of overgrazing. Generality of this regime-shift dynamic is explored across 13 rocky reef systems (spanning 11 different regions from both hemispheres) by compiling available survey data (totalling 10 901 quadrats surveyed in situ) plus experimental regime-shift responses (observed during a total of 57 in situ manipulations). The emergent and globally coherent pattern shows urchin grazing to cause a discontinuous ‘catastrophic’ regime shift, with hysteresis effect of approximately one order of magnitude in urchin biomass between critical thresholds of overgrazing and recovery. Different life-history traits appear to create asymmetry in the pace of overgrazing versus recovery. Once shifted, strong feedback mechanisms provide resilience for each alternative state thus defining the catastrophic nature of this regime shift. Importantly, human-derived stressors can act to erode resilience of desirable macroalgal beds while strengthening resilience of urchin barrens, thus exacerbating the risk, spatial extent and irreversibility of an unwanted regime shift for marine ecosystems.

Background and aims A long-term monitoring program (ranging from 16 to 77 years) on the Dutch Wadden Sea Islands provided well-documented examples of vegetationsuccession in wet dune slacks.We used this opportunity to study soil organic matter (SOM) accumulation in relation to vegetation succession. The aim of this paper is toidentify the factors which regulate accumulation rates of SOM in wet dune slacks.Methods We used several soil chronosequences using data from the monitoring program together with data from a long-term research activity and more recent measurements. We used several soil chronosequences using data from themonitoring program together with data from a long-term research (up to 150 years) and more recent measurements.Fieldmeasurements included pH, soil organicmatter, above ground standing crop and water levels.Water level regimes (inundation duration and mean minimum water level), were simulated using a hydrological model. Capable of simulating inundation duration and water-level fluctuations, thismodel used field measurements collected over more than 5 years, as well as precipitation and evapotranspiration data collected over a period of 25 years.Results Sampling two synchronic chronosequences showed that SOM accumulations increased linearly during the first 50-60 years and then levelled off. Sampling various diachronic chronosequences over time showed a wide variation in accumulation rates. Slacks with low productive species, such as Littorella uniflora, showed low accumulation rates (0.02-0.08 kg/m2/year), and persisted even over a period of more than 90 years. In contrast, slacks dominated by high productive species, such as Phragmites australis, showed ten times higher accumulation rates (0.17-0.26 kg/m2/year) over a similar time period and comparable annual inundation periods (176-240 days). A multiple linear regression showed that variation in SOM accumulation rates was best explained by above-ground biomass of thevegetation. Conclusions We conclude that the rate of SOM accumulationin wet dune slacks is primarily controlled byplant above-ground biomass. Both above-ground biomass and SOM accumulation can remain very low over a long period of time when dune slacks are flooded during most of the year and plants with adaptive traits are able to maintain vegetation succession at a pioneer stage.

Ponds and shallow lakes can be very clear with abundant submerged plants, or very turbid due to a high concentration of phytoplankton and suspended sediment particles. These strongly contrasting ecosystem states have been found to represent alternative attractors with distinct stabilizing feedback mechanisms. In the turbid state, the development of submerged vegetation is prevented by low underwater light levels. The unprotected sediment frequently is resuspended by wave action and by fish searching for food causing a further decrease of transparency. Since there are no plants that could serve as refuges, zooplankton is grazed down by fish to densities insufficient to control algal blooms. In contrast, the clear state in eutrophic shallow lakes is dominated by aquatic macrophytes. The submerged macrophytes prevent sediment resuspension, take up nutrients from the water, and provide a refuge for zooplankton against fish predation. These processes buffer the impacts of increased nutrient loads until they become too high. Consequently, the response of shallow lakes to eutrophication tends to be catastrophic rather than smooth, and various lakes switch back and forth abruptly between a clear and a turbid state repeatedly without obvious external forcing. Importantly, a switch from a turbid to a stable clear state often can be invoked by means of biomanipulation in the form of a temporary reduction of the fish stock.

Changes in trophic status of shallow lakes from the southeastern Pampa plain of Argentina are evaluated based on the relative presence of submerged macrophytes and phytoplankton. The evolution of lake Lonkoy is reconstructed for the last 5000 years, using pollen, non-pollen palynomorphs (NPPs) and macrofossil remain analyses, revealing three main lake stages. A macrophyte-dominated clear lake prevailed until 670 cal yr BP. Afterwards, the lake changed to a turbid phytoplankton-dominated state. Recently, the lake has turned to a turbid phase, caused by resuspended sediment. These results are compared with two other paleolimnological records from the region, lakes Hinojales and Nahuel Rucá, revealing a similar late-Holocene evolution, with a shift from a macrophyte- to phytoplankton-dominated lake, occurring at broadly the same time. The shift of states may be attributed to a shift in climate after 1000 cal. yr BP towards wetter conditions during the Medieval Climate Anomaly.

The traditional perception of wetlands as nutrient sinks has led them to be used as wastewater disposal areas for a long-time, resulting in a severe alteration of the structure and function by eutrophication. Nutrient loading is usually linked to hydrological alterations which encompasses shifts in vegetation patterns and nutrient cycling. The eutrophication process in wetlands accelerates primary productivity and increases net accumulation of organic matter and nutrients but also enhances organic matter decomposition, microbial activity and soluble nutrients in sediments. Internal loading becomes the main nutrient source to the wetland, even in the years of low external inputs, controlling the nutrient dynamics. Since soil phosphorus microbial biomass responds positively to phosphorus enrichment in wetlands, mineralized phosphorus in wetland soils appear as the most responsive microbial indicator to nutrient enrichment in wetlands. Therefore, phosphorus internal loading is the critical factor in regulating eutrophication status of wetlands. N2O and N2 emissions by wetlands can be enhanced in the future as nitrate availability in wetlands continues to be high due to increased pollution

P>The arrival order of colonists in developing populations can have a lasting influence on community and population structure, a phenomenon referred to as priority effects. To explore whether such priority effects are important in determining strain composition of populations of the cyanobacterium Microcystis, four Microcystis strains, isolated from a single lake and differing in functional traits, were grown during 4 weeks in the laboratory in all possible pairwise combinations, with the two strains either inoculated at the same time or with a time lag of 1 week, in the presence or absence of grazing Daphnia magna. The relative abundance of strains in the mixtures was assessed using denaturing gradient gel electrophoresis, and the growth rate of each strain in the mixtures was determined for the last 2 weeks of the experiment. We observed strong effects of inoculation order on the final population structure, and these effects were influenced by grazing Daphnia. The priority effects were strain-specific and occurred in two directions: some of the strains grew slower while others grew faster when inoculated second compared with when inoculated first. Our results indicate that priority effects may have a profound impact on strain composition of Microcystis populations. ispartof: Environmental Microbiology vol:11 issue:10 pages:2564-2573 ispartof: location:England status: published

Droughts can have serious ecological and economic consequences and will pose an increasing challenge to rangeland users as the global climate is changing. Finding ways to reduce ecological and economic impacts of drought should thus be a major research thrust. Resilience, defined as the amount of perturbation a social or ecological system can absorb without shifting to a qualitatively different state, has emerged as a prominent concept in ecosystem ecology and more recently as a conceptual framework for understanding and managing complex social-ecological systems. This paper discusses the application and relevance of resilience to understanding and managing ecosystem change, and enhancing the capacity of land users to adapt to droughts. Drought can trigger vegetation change and factors such as grazing management can influence the likelihood of such transitions. Drought can cause differential mortality of perennial plants and this could provide an opportunity for rangeland restoration by opening up establishment sites for desirable species. The capacity of land users to cope with drought is influenced by the resilience of their agro-ecosystems, the diversity of livelihood options, access to resources and institutional support. By these criteria, current agricultural development approaches in South Africa, particularly in communal rangelands and areas of land reform, are unlikely to enhance land users' resilience to drought and other perturbations.

1. Results are analysed from 11 experiments in which effects of fish addition and nutrient loading on shallow lakes were studied in mesocosms. The experiments, five in 1998, six in 1999, were carried out in six lakes, distributed from Finland to southern Spain, according to a standard protocol. 2. Effects of the treatments on 29 standard chemical, phytoplankton and zooplankton variables are examined to assess the relative importance of bottom-up (nutrient enrichment) and top-down (fish predation) effects. For each year, the experiments in different locations are treated as replicates in a meta-analysis. Results of individual experiments are then compared in terms of the patterns of significant influences of nutrient addition and fish predation with these overall results (the baseline), 3. The overall meta-analysis gave consistent results across the 2 years, with nutrient loading influencing all of the chemical variables, and on average 31% of primary producer and 39% of zooplankton variables. In contrast, fish influenced none of the chemical variables, 11% of the primary producer and 44% of the zooplankton variables. Nutrient effects on the system were thus about three times greater than fish effects, although fish effects were not inconsiderable. 4. The relative importance of nutrients and fish in individual experiments often differed between years at the same location and effects deviated to varying degrees from the baseline. These deviations were treated as measures of consistency (predictability) of conclusions in repeat experiments. Consistency increased southwards and this is interpreted as a consequence of more variable annual weather northwards. 5. The influence of nutrient loading was greater southwards and this was probably manifested through naturally greater annual macrophyte abundance in warmer locations in consequence of the longer plant growing-season. There was no trend in the relative importance of fish effects with latitude but this may partly be an artefact of the simple fish community used. These findings suggest that nutrient control should be a greater priority than biomanipulation in the restoration of eutrophicated shall 6. Starting conditions affected the outcome of experiments. High initial concentrations of total phosphorus and planktonic chlorophyll a concentration (created by local conditions prior to the experiment) led to de-emphasis of the importance of nutrient loading in the experiment. [KEYWORDS: continental gradient ; fish ; meta-analysis ; nutrients ; weather variation]

"This essay discusses three potential models relating disturbance severity to post-disturbance stand composition in the boreal forest: (1) continuous, where changes in disturbance severity cause a proportional and continuous change in stand composition; (2) discontinuous, where a threshold disturbance severity exists beyond which major changes in composition occur; and (3) the cusp, where thresholds exist and coexistence of two alternative compositional states is possible at the same disturbance severity. Any of the three models may be appropriate in different stands or different parts of the boreal forest. If the actual model does not correspond to that assumed by forest managers, then forest harvesting practices may cause unexpected sudden changes in forest composition. Disturbance severity under the natural disturbance regime changes so dramatically, from one disturbance to the next, that oscillations in composition over time are likely to be individualistic and irregular, rather than stable. Harvesting operations are mainly done during winter, resulting in a very low disturbance severity in comparison to the natural regime, which included crown fires. Major changes in species compositon over large parts of the boreal forest are likely to result."

The dynamics of shallow-water coastal environments are controlled by external drivers—sea level rise, storms, and sediment and nutrient supplies—and by internal feedbacks. Interactions of biotic processes (vegetation growth, trophic dynamics) and abiotic drivers can lead to nonlinear responses to changing conditions and to the emergence of thresholds, hysteresis, and alternative stable states. We develop a conceptual framework for studying interactions between the dynamics of marshes and habitats in shallow coastal bays with unconsolidated sediments (seagrass, oyster reefs). Using examples primarily from the Virginia Coast Reserve Long Term Ecological Research site, we show that in the subtidal part of the landscape, two alternative stable states can exist—one dominated by seagrass up to a certain depth that represents a tipping point to the second, unvegetated stable state. The depth limit of the seagrass stable state is influenced by (1) the positive feedback of vegetation on reducing sediment suspension and improving the light environment for growth, (2) climate (e.g., temperature), and (3) water quality. Two stable states are also present in intertidal areas, with salt marshes lying above mean sea level and tidal flats below mean sea level. State transitions are driven by sediment availability, sea level rise, the relative strength of wind waves with respect to tidal currents, and the biotic feedback of vegetation on sediment stabilization and accretion. State-change dynamics in one system may propagate to adjacent systems, and this coupling may influence the landscape-scale response to environmental change. Seagrass meadows and oyster reefs affect adjacent marshes both positively (wave attenuation) and negatively (reduced sediment supply), and marsh-edge erosion could negatively influence the light environment for seagrass growth. Forecasting the resilience of coastal ecosystems and the landscape-scale response to environmental change in the next century requires an understanding of nonlinear dynamics, including the possibility of multiple stable states, the coupled evolution of adjacent systems, and potential early warning signs of thresholds of change.

Background: In European mountains, where semi-natural grasslands have high cultural and nature conservation value, land-use legacies from past ploughing are associated with distinct floristic compositions and ecosystem properties. It remains unknown if these differences are stable or if post-arable grasslands are only transient states that eventually follow a predictable successional pathway. Aims: We use a generic vegetation dynamics model based on plant traits to test these alternative hypotheses in subalpine grasslands of the central French Alps. Methods: We classified dominant graminoids into trait-based Plant Functional Types (PFTs) and simulated their long-term response to fertility, annual mowing and past ploughing. Results: Simulations demonstrated that a temporary dispersal limitation of long-lived tussock grasses could have allowed species-rich hay meadows to develop as alternative communities. We also show that when they are regularly mown, meadows are resistant to colonisation by long-lived tussock grasses. Conclusions: Our results thus support traditional management based on regular mowing, but also highlight the key role of landscape history through its effects on dispersal. Our study also demonstrates the usefulness of easy-to-measure plant traits and a widely applicable generic modelling framework for testing hypotheses on the long-term interactions of multiple drivers of vegetation dynamics.