Search

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).

Multichannel food webs are shaped by the ability of apex predators to link asymmetric energy flows in mesohabitats differing in productivity and community traits. While body size is a fundamental trait underlying life histories and demography, its implications for structuring multichannel food webs are unexplored. To fill this gap, we develop a model that links population responses to predation, and resource availability to community-level patterns, using a tri-trophic food web model with two populations of intermediate consumers and a size-selective top predator. We show that asymmetries in mesohabitat productivities and consumer body sizes drive food web structure, merging previously separate theory on apparent competition and emergent Allee effects (i.e. abrupt population collapses) of top predators. Our results yield theoretical support for empirically observed stability of asymmetric multichannel food webs and discover three novel types of emergent Allee effects involving intermediate consumers, multiple populations or multiple alternative stable states.

Hysteresis is a fundamental characteristic of alternative stable state theory, yet evidence of hysteresis is rare. In mesic grasslands, fire frequency regulates transition from grass- to shrub-dominated system states. It is uncertain, however, if increasing fire frequency can reverse shrub expansion, or if grass-shrub dynamics exhibit hysteresis. We implemented annual burning in two infrequently burned grasslands and ceased burning in two grasslands burned annually. With annual fires, grassland composition converged on that of long-term annually burned vegetation due to rapid recovery of grass cover, although shrubs persisted. When annual burning ceased, shrub cover increased, but community composition did not converge with a long-term infrequently burned reference site because of stochastic and lagged dispersal by shrubs, reflecting hysteresis. Our results demonstrated that annual burning can slow, but not reverse, shrub encroachment. In addition, reversing fire frequencies resulted in hysteresis because vegetation trajectories from grassland to shrubland differed from those of shrubland to grassland.

In mosaic ecosystems, multiple land types coexist as alternative stable states exhibiting distinct spatial patterns. Forest-grassland mosaics are ecologically valuable, due to their high species richness. However, anthropogenic disturbances threaten these ecosystems. Designating protected areas is one approach to preserving natural mosaics. Such work must account for climate change, yet there are few spatially explicit models of mosaics under climate change that can predict its effects. We construct a spatially explicit simulation model for a natural forest-grassland mosaic, parameterized for Southern Brazil. Using this model, we investigate how the spatial structure of these systems is altered under climate change and other disturbance regimes. By including local spatial interactions and fire-mediated forest recruitment, our model reproduces important spatial features of protected real-world mosaics, including the number of forest patches and overall forest cover. Multiple concurrent changes in environmental conditions have greater impacts on tree cover and spatial structure in simulated mosaics than single changes. This sensitivity reflects the narrow range of conditions under which simulated mosaics persist and emphasizes their vulnerability. Our model predicts that, in protected mosaics, climate change impacts on the fire-mediated threshold to recruitment will likely result in substantial increases in forest cover under Representative Concentration Pathway (RCP) 8.5, with potential for mosaic loss over a broad range of initial forest cover levels. Forest cover trajectories are similar until 2150, when cover increases under RCP 8.5 outpace those under RCP 2.6. Mosaics that persist under RCP 8.5 may experience structural alterations at the patch and landscape level. Our simple model predicts several realistic aspects of spatial structure as well as plausible responses to likely regional climate shifts. Hence, further model development could provide a useful tool when building strategies for protecting these ecosystems, by informing site selection for conservation areas that will be favourable to forest-grassland mosaics under future climates.

Mutualistic interactions between species are ubiquitous in nature and essential for ecosystem functioning. Often dozens or even hundreds of species with different degrees of specialisation form complex networks. How this complexity evolves is a fundamental question in ecology. Here, we present a new game theoretical approach to model complex coevolutionary processes and apply it to pollination networks. A theoretical analysis reveals multiple evolutionary stable network structures that depend on the availability of pollination service. In particular, we find efficient communities, in which a high percentage of pollen are transported conspecifically, to evolve only when plant and pollinator abundances are well balanced. Both pollinator shortage and oversupply select for more inefficient network structures. The results suggest that availability of pollination services is a key factor structuring pollination networks and may offer a new explanation for geographical differences in pollination communities that have long been recognised by ecologists.

As a result of climate and land‐use changes, grasslands have been subjected to intensifying drought regimes. Extreme droughts could interfere in the positive feedbacks between grasses and soil water content, pushing grasslands across critical thresholds of productivity and leading them to collapse. If this happens, systems may show hysteresis and costly management interventions might be necessary to restore predrought productivity. Thus, neglecting critical transitions may lead to mismanagement of grasslands and to irreversible loss of ecosystem services. Rainfall manipulation experiments constitute a powerful approach to investigate the risk of such critical transitions. However, experiments performed to date have rarely applied extreme droughts and have used resilience indices that disregard the existence of hysteresis. Here, we suggest how to incorporate critical transitions when designing rainfall manipulation experiments on grasslands and when measuring their resilience to drought. The ideas presented here have the potential to trigger a perspective shift among experimental researchers, into a new state where the existence of critical transitions will be discussed, experimentally tested, and largely considered when assessing and managing vegetation resilience to global changes., We suggest two different approaches of how rainfall manipulation experiments in grasslands can be carried out to investigate critical transitions of productivity. We aim to trigger a perspective shift and guide researchers along the trajectory to a new state, where the existence of critical transitions will be discussed, experimentally tested, and largely considered when assessing and managing vegetation resilience to global changes.

There is growing acknowledgement that human���induced change can push ecosystems beyond tipping points, resulting in the dramatic and sudden loss of vital ecosystem services. Invasive non���native species (INNS) are spreading rapidly due to anthropogenic activities and climate change and can drive changes to ecosystem functioning by altering abiotic conditions and restructuring native communities. Shallow lake ecosystems are especially vulnerable to perturbation from INNS as they can exist in two alternative stable states: either clear water with an abundance of vegetation or turbid, unvegetated and dominated by phytoplankton. Through a global meta���analysis of studies observing the effects of INNS on recipient lake ecosystems, we found that certain INNS drive significant changes in the abundance of key taxa and conditions that govern the balance of alternative equilibria in shallow lakes. Invasive fish and crustaceans demonstrated effects likely to lead to early ecosystem collapse to a turbid state and delay ecosystem recovery. Invasive molluscs presented opposite effects, which may delay ecosystem collapse and encourage ecosystem recovery. Our results demonstrate that INNS could significantly alter the tipping points of ecosystem collapse and recovery, and that not all invasive species may initiate system collapse. Our results provide guidance for managers of invaded shallow lake ecosystems, which provide diverse services including sanitation, potable water supply, industrial cooling, aquaculture and recreational resources. Moreover, our approach could be applied to identify key potential drivers of change in other crucial ecosystems which demonstrate alternative equilibria, such as coral reefs and kelp forests.

Many arid and semi‐arid rangelands exhibit distinct spatial patterning of vegetated and bare soil‐dominated patches. The latter potentially represent a grazing‐induced, degraded ecosystem state, but could also arise via mechanisms related to feedbacks between vegetation cover and soil moisture availability that are unrelated to grazing. The degree to which grazing contributes to the formation or maintenance of degraded patches has been widely discussed and modeled, but empirical studies of the role of grazing in their formation, persistence, and reversibility are limited.We report on a long‐term (17 years) grazing removal experiment in a semi‐arid savanna where vegetated patches composed of perennial grasses were interspersed within large (>10 m2) patches of bare soil.Short‐term (3 years) grazing removal did not allow bare patches to become revegetated, whereas following long‐term (17 years) grazing removal, bare soil patches were revegetated by a combination of stoloniferous grasses and tufted bunchgrasses. In the presence of grazers, stoloniferous grasses partially recolonized bare patches, but this did not lead to full recovery or to the establishment of tufted bunchgrasses.These results show that grazers alter both the balance between bare and vegetated patches, as well as the types of grasses dominating both patch types in this semiarid savanna.Synthesis: Large herbivores fundamentally shaped the composition and spatial pattern of the herbaceous layer by maintaining a two‐phase herbaceous mosaic. However, bare patches within this mosaic can recover given herbivore removal over sufficiently long time scales, and hence do not represent a permanently degraded ecosystem state., The stability and spatial pattern of degraded and vegetated patches in dryland ecosystems has been modeled and discussed for decades, with few empirical studies. We present one of the first long‐term experiments that demonstrate these patterns are maintained by large herbivore grazing and can disappear when herbivores are removed.

AIM: Previous analyses of remotely sensed data detected the multimodality of the tree‐cover distribution of the boreal forest, and identified areas with potentially alternative tree‐cover states. This paper aims at investigating the causes of multimodality and multistability of the boreal forest, their influence on the asymmetric tree species distribution between Eurasia and North America, and whether multistability could be associated with recent greening trends in leaf area index (LAI) and normalized difference vegetation index (NDVI). LOCATION: Eurasian and North American boreal forests. TIME PERIOD: 2000–2010. MAJOR TAXA STUDIED: Boreal forest plant functional types. METHODS: We employ a conceptual model based on tree species competition to simulate the sensitivity of tree cover to stochastic disturbances and to changes in environmental factors. We include different plant functional types based on survival adaptations, and force the model with remotely sensed environmental data. We analyse the model as a dynamical system. We use metrics from statistics and information theory to compare the detection of alternative tree‐cover states and greening trends in LAI and NDVI. RESULTS: We find that multimodality and multistability can emerge through competition between different plant functional types. Additionally, our model is able to reproduce the asymmetry in tree species distribution between Eurasia and North America. Moreover, changes in permafrost distribution can be associated with phenomenological bifurcation points of the model. Finally, we find that the detection of multistable areas is not affected by recent vegetation trends, whereas shifts between alternative states could have affected the greening trends. MAIN CONCLUSIONS: Tree‐cover multistability in the boreal region can emerge through competition between species subject to periodic disturbances. Changes in permafrost thaw and distribution could be responsible for the asymmetry in tree species distribution between North America and Eurasia. Climate change and permafrost degradation could cause shifts in tree‐cover states and dominant species. Recent vegetation greening trends in multistable areas could have been affected by shifts between alternative states.

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.

Predation can affect prey behavior, demography, abundance, and distribution, particularly in lentic freshwater ecosystems. Fish are predators known to reduce the abundance of their prey and to restrict the distribution of species. Using time series which spanned 43 and 22 yr, respectively, we analyzed the effect of a change in the fish predator community on the dynamics of two pond-breeding amphibian populations (Rana temporaria and Rana dalmatina). Specifically, we used a state-space time series model which allows for density dependence and observation error, to ask whether the change in predation risk affects population growth rate and the return point around which the populations fluctuate. The results showed that the type of observation error assumed did not affect the biological parameters. We found evidence for density dependence in both populations. The effect of the change in fish predation on population growth rate and the return point was strong in the population where fish invaded a previously fish-free pond. The effect was weaker in the population where the change was from cyprinid fish to pike. The results showed that fish predation can have strong effects on amphibian population dynamics. The observed population dynamical pattern is phenomenologically similar to alternative stable states.

QUESTIONS: Models predicting the distribution of savannas worldwide have identified rainfall and fire as their primary determinants. However, most of them have relied upon adult traits, while juvenile traits, at the bottleneck of the plant's life cycle, have been largely overlooked. We developed a novel mechanistic model based on the effects of temperature and fire on germination traits to predict the distribution of Cerrado, i.e., the largest neotropical savanna. LOCATION: Cerrado and neighboring biomes. METHODS: We compiled data on the germination of seeds subjected to temperature and heat shock treatments and used generalized additive mixed models to predict germination potential as a function of temperature, species, physiognomy (forest/savanna), habits (herbs/shrubs/trees), and fire frequency. RESULTS: The best model showed that seasonal temperatures set the germination limits for seeds of both savanna and forest physiognomies. Forest seeds presented a higher germinability in the optimum temperature range, but savanna seeds had higher survival rates after heat shocks. The model revealed that the southern limit of Cerrado is determined by low winter temperatures, while the western and eastern boundaries are set by high summer temperatures. The model also predicted an area of high germination potential that coincides with high biodiversity and climate stability in the Cerrado. CONCLUSIONS: Germination traits are highly valuable to predict vegetation responses to climate. Seasonal temperatures are primary determinants of the Cerrado’s extent, while fire favors the recruitment of savanna species over the Cerrado–Amazonia ecotone. Global warming may significantly impact the germination potential of native species.

QUESTIONS: Increased wildfire activity is resulting in plant community‐type conversions worldwide. In some regions, fire‐sensitive forests are being replaced by flammable fire‐resilient communities, increasing the likelihood of reburning due to positive fire feedbacks. Here we evaluated whether fire severity affects post‐fire plant community flammability attributes that lead to community‐type conversions and changes in the likelihood of reburning. Specifically, we assessed how fire severity, the dominant pre‐fire vegetation, and distance to unburned remnant forest may shape post‐fire community properties and flammability trajectories in northwestern Patagonia. LOCATION: Northwestern Patagonia, Argentina. METHODS: We repeated sampling of Nothofagus pumilio, Nothofagus dombeyi, and Austrocedrus chilensis forests and native shrublands (composed of multiple shrub species) two and 18 years after fires that burned at low‐ to very high‐severity levels, and measured community structure and flammability attributes. RESULTS: Eighteen years after fire, forests that burned at moderate to very high severity were unable to recover and were replaced by more flammable shrublands and grasslands. Following low‐severity fire, fine‐fuel density was lower and forest recovery was enhanced by greater survival of remnant seed trees. Burnt shrublands increased in abundance across all severity classes but attained highest fine‐fuel production after moderate‐ to very high‐severity fire. CONCLUSIONS: Low fire severity, by enabling regeneration of forests that are less flammable than shrublands, diminishes the likelihood of reburning, thus establishing a negative feedback favorable to forest perpetuation. Conversely, moderate to very high fire severity leads to a positive feedback by promoting conversion to shrublands and greater fine‐fuel accumulation. This increases the likelihood of reburning, reinforcing the persistence of pyrophytic communities and favoring landscape‐scale loss of fire‐sensitive forests. This fire severity‐mediated positive feedback may in many regions worldwide further amplify warming‐related wildfire activity increases, posing serious threats to the persistence of fire‐sensitive ecosystems.

Ecologists have proposed that montane grassland-shola (stunted evergreen forest) mosaics in the Western Ghats may represent alternative stable vegetation states. But paleoecology investigations sel...

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.

Shifting disturbance regimes can have cascading effects on many ecosystems processes. This is particularly true when the scale of the disturbance no longer matches the regeneration strategy of the dominant vegetation. In the yellow pine and mixed conifer forests of California, over a century of fire exclusion and the warming climate are increasing the incidence and extent of stand-replacing wildfire; such changes in severity patterns are altering regeneration dynamics by dramatically increasing the distance from live tree seed sources. This has raised concerns about limitations to natural reforestation and the potential for conversion to non-forested vegetation types, which in turn has implications for shifts in many ecological processes and ecosystem services. We used a California region-wide data set with 1,848 plots across 24 wildfires in yellow pine and mixed conifer forests to build a spatially explicit habitat suitability model for forecasting postfire forest regeneration. To model the effect of seed availability, the critical initial biological filter for regeneration, we used a novel approach to predicting spatial patterns of seed availability by estimating annual seed production from existing basal area and burn severity maps. The probability of observing any conifer seedling in a 60-m2 area (the field plot scale) was highly dependent on 30-yr average annual precipitation, burn severity, and seed availability. We then used this model to predict regeneration probabilities across the entire extent of a "new" fire (the 2014 King Fire), which highlights the spatial variability inherent in postfire regeneration patterns. Such forecasts of postfire regeneration patterns are of importance to land managers and conservationists interested in maintaining forest cover on the landscape. Our tool can also help anticipate shifts in ecosystem properties, supporting researchers interested in investigating questions surrounding alternative stable states, and the interaction of altered disturbance regimes and the changing climate.

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.

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.

Early warning signals (EWS) are statistical indicators that a rapid regime shift may be forthcoming. Their development has given ecologists hope of predicting rapid regime shifts before they occur. Accurate predictions, however, rely on the signals being appropriate to the system in question. Most of the EWS commonly applied in ecology have been studied in the context of one specific type of regime shift (the type brought on by a saddle-node bifurcation, at which one stable equilibrium point collides with an unstable equilibrium and disappears) under one particular perturbation scheme (temporally uncorrelated noise that perturbs the net population growth rate in a density independent way). Whether and when these EWS can be applied to other ecological situations remains relatively unknown, and certainly underappreciated. We study a range of models with different types of dynamical transitions (including rapid regime shifts) and several perturbation schemes (density-dependent uncorrelated or temporally-correlated noise) and test the ability of EWS to warn of an approaching transition. We also test the sensitivity of our results to the amount of available pre-transition data and various decisions that must be made in the analysis (i.e. the rolling window size and smoothing bandwidth used to compute the EWS). We find that EWS generally work well to signal an impending saddle-node bifurcation, regardless of the autocorrelation or intensity of the noise. However, EWS do not reliably appear as expected for other types of transition. EWS were often very sensitive to the length of the pre-transition time series analyzed, and usually less sensitive to other decisions. We conclude that the EWS perform well for saddle-node bifurcation in a range of noise environments, but different methods should be used to predict other types of regime shifts. As a consequence, knowledge of the mechanism behind a possible regime shift is needed before EWS can be used to predict it.

1.Biotic homogenization (BH), a process by which β-diversity erodes, represents a severe threat to biodiversity. Wetland plant communities may be especially susceptible to BH; however, this process has rarely been quantified and represents just one of many possible outcomes of compositional change. Likewise, few studies have used more than two timepoints to investigate BH as a dynamic temporal process. 2.To address these issues, we quantified changes in β-diversity amongst 48 herbaceous emergent wetlands across a landscape over four timepoints from 1997 to 2015. Pairwise occurrence- and abundance-based dissimilarity metrics were used to quantify β-diversity. Nonmetric multidimensional scaling was used to map these changes in compositional similarity through time, and repeated measures ANOVA via randomization was used to test for a significant effect of time on β-diversity. 3.Results indicated that herbaceous emergent wetlands homogenized in Illinois over 15 years due to the increased presence and abundance of Phalaris arundinacea and the decline of several other species. Temporal dynamics of β-diversity differed between occurrence- and abundance-based analyses. 4.Synthesis Using both presence-absence and abundance data to investigate BH helped avoid underestimating the impact of BH on β-diversity, and drew attention to different temporal dynamics in β-diversity at these two resolutions of community structure. This study highlights challenges associated with investigating BH, such as problems documenting the process where homogenization is not occurring in every site in a region, or where sites are converging towards more than one homogenized state. Incorporating concepts like alternative stable states may be helpful in resolving these issues, and present a more realistic picture of ecological change. This article is protected by copyright. All rights reserved.

© 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.

Research on lake eutrophication often identifies variables affecting amounts of phosphorus (P) and nitrogen (N) in lakes, but understanding factors influencing N:P ratios is important given its influence on species composition and toxin production by cyanobacteria. We sampled 80 shallow lakes in Minnesota (USA) for three years to assess effects of watershed size, proportion of watershed as both row crop and natural area, fish biomass, and lake alternative state (turbid vs. clear) on total N : total P (TN : TP), ammonium, total dissolved phosphorus (TDP), and seston stoichiometry. We also examined N:P stoichiometry in 20 additional lakes that shifted states during the study. Last, we assessed the importance of denitrification by measuring denitrification rates in sediment cores from a subset of 34 lakes, and by measuring seston δ15 N in four additional experimental lakes before and after they were experimentally manipulated from turbid to clear states. Results showed alternative state had the largest influence on overall N:P stoichiometry in these systems, as it had the strongest relationship with TN : TP, seston C:N:P, ammonium, and TDP. Turbid lakes had higher N at given levels of P than clear lakes, with TN and ammonium 2-fold and 1.4-fold higher in turbid lakes, respectively. In lakes that shifted states, TN was 3-fold higher in turbid lakes, while TP was only 2-fold higher, supporting the notion N is more responsive to state shifts than is P. Seston δ15 N increased after lakes shifted to clear states, suggesting higher denitrification rates may be important for reducing N levels in clear states, and potential denitrification rates in sediment cores were among the highest recorded in the literature. Overall, our results indicate lake state was a primary driver of N:P dynamics in shallow lakes, and lakes in clear states had much lower N at a given level of P relative to turbid lakes, likely due to higher denitrification rates. Shallow lakes are often managed for the clear-water state due to increased value as wildlife habitat. However, our results indicate lake state also influences N biogeochemistry, such that managing shallow lakes for the clear-water state may also mitigate excess N levels at a landscape scale.

As ecology and evolution become ever more entwined, many areas of ecological theory are being re-examined. Eco-evolutionary analyses of classic coexistence mechanisms are yielding new insights into the structure and stability of communities. We examine fluctuation-dependent coexistence models, identifying communities that are both ecologically and evolutionarily stable. Members of these communities possess distinct environmental preferences, revealing widespread patterns of limiting similarity. This regularity leads to consistent changes in the structure of communities across fluctuation regimes. However, at high amplitudes, subtle differences in the form of fluctuations dramatically affect the collapse of communities. We also show that identical fluctuations can support multiple evolutionarily stable communities - a novel example of alternative stable states within eco-evolutionary systems. Consequently, the configuration of communities will depend on historical contingencies, including details of the adaptive process. Integrating evolution into the study of coexistence offers new insights, while enriching our understanding of ecology.

Ecological memory, often determined by the extent and type of retained biological legacies present following disturbance, may produce persistent landscape patterns. However, after fire, the persistence or switch to an alternative state may depend on the complex interplay of ecological memory (biological legacies) and potential effects of new external factors influencing the post-fire environment. The current study assesses both the strength of ecological memory resulting from biological legacies of pre-burn vegetation types as well as post-fire effects of livestock. Following a severe fire in 1999, we set up a network of long-term exclosures to examine the effects of legacies and cumulative herbivory by cattle on fuel types, amounts, distribution, flammability and microenvironmental conditions in two post-fire communities representing alternative fire-driven states: pyrophobic Nothofagus pumilio subalpine forests and pyrophytic Nothofagus antarctica tall shrublands in northwestern Patagonia, Argentina. Our results show that the retained post-disturbance legacies of tall shrublands and subalpine forests largely determine fuel and flammability traits of the post-fire plant communities 16 years after fire. The importance of biological legacies retained from the unburned plant communities was reflected by the substantially higher amounts of total fine fuel, greater vertical and horizontal fuel continuity and the higher temperatures reached during experimental tissue combustion at post-fire shrubland compared to post-fire forest sites. We show that herbivores may produce antagonistic effects on flammability by decreasing tissue ignitability, total fine fuel and litter depth, and disrupting the vertical and horizontal fine fuel continuity, therefore reducing the probability of fire propagation. However, cattle can increase ratios of dead to live fine fuels, reduce soil moisture, and inhibit tree height growth to canopy size, consequently impeding the development of a closed pyrophobic forest canopy. Synthesis. Our results support the hypothesis that biological legacies, most importantly the dominance by pyrophytic woody plants that resprout vigorously vs. the dominance by pyrophobic obligate seeders, favour fuel and flammability characteristics at the community level which reinforce the mechanisms maintaining pyrophytic shrublands vs. pyrophobic forests. Herbivory by introduced cattle can partially blur sharp pyrophobic/pyrophytic state boundaries by promoting the development of novel post-fire transitional states. Fil: Blackhall, Melisa. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: Raffaele, Estela. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: Paritsis, Juan. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: Tiribelli, Florencia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: Morales, Juan Manuel. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: Kitzberger, Thomas. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: Gowda, Juan Janakiram Haridas. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina Fil: Veblen, Thomas. University of Colorado; Estados Unidos

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.