Species packing in eco‐evolutionary models of seasonally fluctuating environments

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.     

长江口滨海湿地潮间带生态系统的多稳态特征

多稳态现象普遍存在于多种生态系统中,它与生态系统的健康和可持续发展密切相关,已成为生态学研究的热点与难点,但是目前有关滨海湿地生态系统多稳态的形成机制还缺乏深入研究.本文以崇明东滩鸟类自然保护区的潮间带生态系统为研究对象,通过以下内容,开展滨海湿地多稳态研究: 1)通过验证多稳态的判定依据“双峰”和“阈值”特征,证实长江口潮间带生态系统存在多稳态,并确定其稳态类型;2)通过监测潮间带生态系统水动力过程、沉积动力过程以及盐沼植物生长和扩散情况,分析盐沼植被与沉积地貌之间的正反馈作用,进而探讨潮间带生态系统多稳态的形成机制.结果表明: 1)潮间带生态系统的归一化植被指数(NDVI)频度分布存在明显的双峰特征,且盐沼植物成活存在生物量阈值效应,均证实潮间带生态系统存在多稳态,“盐沼”和“光滩”是潮间带生态系统的两种相对稳定状态;2)崇明东滩盐沼前沿的沉积地貌表现出泥沙快速淤积的趋势,显著促进了盐沼植物的生长,盐沼植物与泥沙淤积之间的这种正反馈作用是潮间带生态系统形成多稳态的主要原因;3)盐沼植被扩散格局监测结果在景观尺度上也表明,泥沙淤积作用促进了潮间带生态系统“盐沼”和“光滩”多稳态的形成.本研究既丰富了滨海湿地稳态转换的机理研究,也为我国开展海岸带保护、修复和管理提供了科学依据,具有重要的理论和实践意义.     

Foreseeing the future of mutualistic communities beyond collapse

Changing conditions may lead to sudden shifts in the state of ecosystems when critical thresholds are passed. Some well‐studied drivers of such transitions lead to predictable outcomes such as a turbid lake or a degraded landscape. Many ecosystems are, however, complex systems of many interacting species. While detecting upcoming transitions in such systems is challenging, predicting what comes after a critical transition is terra incognita altogether. The problem is that complex ecosystems may shift to many different, alternative states. Whether an impending transition has minor, positive or catastrophic effects is thus unclear. Some systems may, however, behave more predictably than others. The dynamics of mutualistic communities can be expected to be relatively simple, because delayed negative feedbacks leading to oscillatory or other complex dynamics are weak. Here, we address the question of whether this relative simplicity allows us to foresee a community's future state. As a case study, we use a model of a bipartite mutualistic network and show that a network's post‐transition state is indicated by the way in which a system recovers from minor disturbances. Similar results obtained with a unipartite model of facilitation suggest that our results are of relevance to a wide range of mutualistic systems.     

Alternative (un)stable states in a stochastic predator–prey model

Stochastic models sometimes behave qualitatively differently from their deterministic analogues. We explore the implications of this in ecosystems that shift suddenly from one state to another. This phenomenon is usually studied through deterministic models with multiple stable equilibria under a single set of conditions, with stability defined through linear stability analysis. However, in stochastic systems, some unstable states can trap stochastic dynamics for long intervals, essentially masquerading as additional stable states. Using a predator–prey model, we demonstrate that this effect is sufficient to make a stochastic system with one stable state exhibit the same characteristics as an analogous system with alternative stable states. Although this result is surprising with respect to how stability is defined by standard analyses, we show that it is well-anticipated by an alternative approach based on the system's “quasi-potential.” Broadly, understanding the risk of sudden state shifts will require a more holistic understanding of stability in stochastic systems.     

Shallow lakes theory revisited: various alternative regimes driven by climate,nutrients, depth and lake size

Shallow lakes have become the archetypical example of ecosystems with alternative stable states. However, since the early conception of that theory, the image of ecosystem stability has been elaborated for shallow lakes far beyond the simple original model. After discussing how spatial heterogeneity and fluctuation of environmental conditions may affect the stability of lakes, we review work demonstrating that the critical nutrient level for lakes to become turbid is higher for smaller lakes, and seems likely to be affected by climatic change too. We then show how the image of just two contrasting states has been elaborated. Different groups of primary producers may dominate shallow lakes, and such states dominated by a particular group may often represent alternative stable states. In tropical lakes, or small stagnant temperate waters, free-floating plants may represent an alternative stable state. Temperate shallow lakes may be dominated alternatively by charophytes, submerged angiosperms, green algae or cyanobacteria. The change of the lake communities along a gradient of eutrophication may therefore be seen as a continuum in which gradual species replacements are interrupted at critical points by more dramatic shifts to a contrasting alternative regime dominated by different species. The originally identified shift between a clear and a turbid state remains one of the more dramatic examples, but is surely not the only discontinuity that can be observed in the response of these ecosystems to environmental change.     

Alternative transient states and slow plant community responses after changed flooding regimes

Climate change will have large consequences for flooding frequencies in freshwater systems. In interaction with anthropogenic activities (flow regulation, channel restoration and catchment land‐use) this will both increase flooding and drought across the world. Like in many other ecosystems facing changed environmental conditions, it remains difficult to predict the rate and trajectory of vegetation responses to changed conditions. Given that critical ecosystem services (e.g. bank stabilization, carbon subsidies to aquatic communities or water purification) depend on riparian vegetation composition, it is important to understand how and how fast riparian vegetation responds to changing flooding regimes. We studied vegetation changes over 19 growing seasons in turfs that were transplanted in a full‐factorial design between three riparian elevations with different flooding frequencies. We found that (a) some transplanted communities may have developed into an alternative stable state and were still different from the target community, and (b) pathways of vegetation change were highly directional but alternative trajectories did occur, (c) changes were rather linear but faster when flooding frequencies increased than when they decreased, and (d) we observed fastest changes in turfs when proxies for mortality and colonization were highest. These results provide rare examples of alternative transient trajectories and stable states under field conditions, which is an important step towards understanding their drivers and their frequency in a changing world.     

Vegetation change and conservation status of Coastal Upland Swamps

Coastal Upland Swamp communities are characterized by high biodiversity and provide habitat for a range of threatened flora and fauna. In this research project, we are monitoring swamp vegetation dynamics over decadal timescales and relating observed changes to environmental factors. We have also modelled potential effects of climate change on swamp distributions. We found that swamp communities are spatially dynamic, both internally and in relation to the woodland matrix. Transitions between communities depended on initial states. In addition, these water‐dependent communities appeared highly sensitive to projected climate change and their ‘Endangered’ status makes their active management a high priority. Improved understanding of dynamics at the community and landscape scale facilitates horizon scanning and improves our capacity to plan effective management interventions now and under future management and climate change scenarios.     

Gallery Forest or Herbaceous Wetland? The Need for Multi‐Target Perspectives in Riparian Restoration Planning

Much riparian restoration focuses on establishment of gallery forests, with relatively limited effort to restore herbaceous wetlands as key components of riparian landscape mosaics. Multiple reasons for this include inherent cultural or esthetic preferences, greater availability of scientific knowledge to support riparian forest restoration, and choices of ecological indicators commonly used for monitoring and assessment. Yet riparian herbaceous wetlands have declined dramatically as a result of river regulation and agricultural development, leading to losses of important habitats and ecosystem services that differ from those provided by gallery forests. As an alternative to a single‐minded focus on tree establishment, we advocate restoration of diverse and dynamic habitat mosaics in the context of natural variability of flow and sediment regimes. Landscape context should inform active restoration activities at the local scale, such that riparian forests are not planted in ecologically inappropriate sites. Models are needed to match life history requirements of particular wetland herbaceous plant species with details of flow and sediment transport regimes. We emphasize the importance of herbaceous wetlands as a critical and often overlooked component of riparian ecosystems, and the need for both passive and active restoration of fluvial marshes, sloughs, wet meadows, alkali meadows, off‐channel ephemeral ponds, and other critical floodplain communities associated with herbaceous plant dominance.     

Mechanical vulnerability explains size‐dependent mortality of reef corals

Understanding life history and demographic variation among species within communities is a central ecological goal. Mortality schedules are especially important in ecosystems where disturbance plays a major role in structuring communities, such as coral reefs. Here, we test whether a trait‐based, mechanistic model of mechanical vulnerability in corals can explain mortality schedules. Specifically, we ask whether species that become increasingly vulnerable to hydrodynamic dislodgment as they grow have bathtub‐shaped mortality curves, whereas species that remain mechanically stable have decreasing mortality rates with size, as predicted by classical life history theory for reef corals. We find that size‐dependent mortality is highly consistent between species with the same growth form and that the shape of size‐dependent mortality for each growth form can be explained by mechanical vulnerability. Our findings highlight the feasibility of predicting assemblage‐scale mortality patterns on coral reefs with trait‐based approaches.     

Scale dependence of temperature as an abiotic driver of species' distributions

Aim Scale dependence of patterns and processes remains one of the major unresolved problems in ecology. The responses of ecosystems to environmental stressors are reported to be strongly scale dependent, but projections of the effects of climate change on species' distributions are still restricted to particular scales and knowledge about scale dependence is lacking. Here we propose that the scale dependence of those species' niche dimensions related to climate change is strongly related to the strength of climatic cross‐scale links. More specifically, we hypothesize that the strong cross‐scale links between micro‐ and macroclimatic conditions are related to high cross‐scale similarity (low scale dependence) of species' realized temperature niches and, thus, species' spatial distributions. Location This study covers seven orders of magnitude of spatial scale, ranging from local‐scale (below a metre) and regional‐scale (kilometre) investigations in central European wetland ecosystems to continental‐scale (thousands of kilometres) studies of species' distributions. Methods We combined data on the spatial occurrence of species (vegetation records at local and regional scales, digitized distribution maps at the continental scale) with information about the corresponding temperature regime of vascular plant species occurring in environmentally stable wetland ecosystems characterized by strong cross‐scale links between micro‐ and macroclimatic conditions. Results We observed high cross‐scale similarity of the characteristics of species temperature niches across seven orders of magnitude of spatial scale. However, the importance of temperature as an abiotic driver decreased nonlinearly with decreasing scale, suggesting greater importance of additional (biotic) drivers of species' occurrence at small spatial scales. Main conclusions We report high cross‐scale similarity of realized temperature niches for species inhabiting ecosystems where small‐scale environmental noise is low and cross‐scale links between micro‐ and macroclimatic conditions are strong. By highlighting a strong relationship between abiotic and biotic cross‐scale similarity, our results will help to improve niche‐based species distribution modelling, one of the major assessment tools for determining the ecological effects of climate change.     

Endocrine disruption in aquatic systems: up‐scaling research to address ecological consequences

Endocrine‐disrupting chemicals (EDCs) can alter biological function in organisms at environmentally relevant concentrations and are a significant threat to aquatic biodiversity, but there is little understanding of exposure consequences for populations, communities and ecosystems. The pervasive nature of EDCs within aquatic environments and their multiple sub‐lethal effects make assessments of their impact especially important but also highly challenging. Herein, we review the data on EDC effects in aquatic systems focusing on studies assessing populations and ecosystems, and including how biotic and abiotic processes may affect, and be affected by, responses to EDCs. Recent research indicates a significant influence of behavioural responses (e.g. enhancing feeding rates), transgenerational effects and trophic cascades in the ecological consequences of EDC exposure. In addition, interactions between EDCs and other chemical, physical and biological factors generate uncertainty in our understanding of the ecological effects of EDCs within aquatic ecosystems. We illustrate how effect thresholds for EDCs generated from individual‐based experimental bioassays of the types commonly applied using chemical test guidelines [e.g. Organisation for Economic Co‐operation and Development (OECD)] may not necessarily reflect the hazards associated with endocrine disruption. We argue that improved risk assessment for EDCs in aquatic ecosystems urgently requires more ecologically oriented research as well as field‐based assessments at population‐, community‐ and food‐web levels.     

Assessing the hydrologic impact of historical railroad embankments on wetland vegetation response in Canaan Valley,West Virginia: the value of high‐resolution data

The recovery of natural ecological processes after disturbance is poorly understood. Some disturbances may be so severe as to set ecosystems onto a new trajectory. The Canaan Valley National Wildlife Refuge in West Virginia protects a unique high‐altitude wetland that was heavily disturbed by logging 100 years before present (BP) and has since transitioned to a new ecological state (shrub wetland). Refuge managers interested in preserving and restoring ecosystem states expressed concern about lingering impacts of previous disturbances (logging, railroads, beaver, deer, fire). Available data suggested hydrologic impacts from a remnant railroad grade, but managers had insufficient quantitative data to assess these impacts. We initiated a fine‐scale assessment of topography, vegetation distribution, and hydrology to assess impacts from the remnant rail grade using lidar data, vegetation surveys, and piezometers. We developed topographic models, hydrological models, and mapped vegetation distribution. We developed statistical models to assess relationships between vegetation communities, hydrology, and distance to the rail grade. Surprisingly, we found that hydrologic flow paths did not conform to expectation and were not restricted by remnant land use features. For the most part, vegetation communities are responding to topographic and environmental gradients that existed prior to disturbance. Use of highly detailed topographic data (lidar), field hydrology, and vegetation studies allowed us to more accurately assess hydrologic and vegetation regimes, eliminating the need for mitigation, thus saving significant resources.     

Resilience and restoration of tropical and subtropical grasslands,savannas, and grassy woodlands

Despite growing recognition of the conservation values of grassy biomes, our understanding of how to maintain and restore biodiverse tropical grasslands (including savannas and open‐canopy grassy woodlands) remains limited. To incorporate grasslands into large‐scale restoration efforts, we synthesised existing ecological knowledge of tropical grassland resilience and approaches to plant community restoration. Tropical grassland plant communities are resilient to, and often dependent on, the endogenous disturbances with which they evolved – frequent fires and native megafaunal herbivory. In stark contrast, tropical grasslands are extremely vulnerable to human‐caused exogenous disturbances, particularly those that alter soils and destroy belowground biomass (e.g. tillage agriculture, surface mining); tropical grassland restoration after severe soil disturbances is expensive and rarely achieves management targets. Where grasslands have been degraded by altered disturbance regimes (e.g. fire exclusion), exotic plant invasions, or afforestation, restoration efforts can recreate vegetation structure (i.e. historical tree density and herbaceous ground cover), but species‐diverse plant communities, including endemic species, are slow to recover. Complicating plant‐community restoration efforts, many tropical grassland species, particularly those that invest in underground storage organs, are difficult to propagate and re‐establish. To guide restoration decisions, we draw on the old‐growth grassland concept, the novel ecosystem concept, and theory regarding tree cover along resource gradients in savannas to propose a conceptual framework that classifies tropical grasslands into three broad ecosystem states. These states are: (1) old‐growth grasslands (i.e. ancient, biodiverse grassy ecosystems), where management should focus on the maintenance of disturbance regimes; (2) hybrid grasslands, where restoration should emphasise a return towards the old‐growth state; and (3) novel ecosystems, where the magnitude of environmental change (i.e. a shift to an alternative ecosystem state) or the socioecological context preclude a return to historical conditions.     

The effect of seasonal variation in abiotic factors on fish community structure in temporary and permanent pools in a tropical wetland

1. Fish community structure depends on biotic interactions and abiotic variables. Abiotic variables appear to gain importance in highly variable freshwater systems, such as tropical wetlands where a marked seasonal hydroperiod (dry and wet seasons) modifies water quality and quantity, differentially affecting fish survival and, consequently, modifying species richness and abundance. 2. We evaluated the relationship between abiotic variables and fish community structure in variable (temporary) and stable (permanent) pools that were interconnected in a tropical wetland with marked annual dry and wet seasons. 3. All fish species were able to occupy any of the studied pools, but our results showed distinctive fish community structures in permanent and temporary pools. Community structure was related to temperature, depth, pH and macrophyte coverage. Total fish abundance in the wetland was negatively related to water depth and positively related to macrophyte coverage. 4. Null models of co‐occurrence indicated a non‐random pattern at the wetland scale and a random pattern within groups of pools with similar characteristics, suggesting that fish communities are structured according to habitat features. We conclude that seasonal abiotic variation and habitat characteristics in this highly variable pristine wetland play major roles in structuring fish communities.     

Community assembly: alternative stable states or alternative transient states?

The concept of alternative stable states has long been a dominant framework for studying the influence of historical contingency in community assembly. This concept focuses on stable states, yet many real communities are kept in a transient state by disturbance, and the utility of predictions for stable states in explaining transient states remains unclear. Using a simple model of plant community assembly, we show that the conditions under which historical contingency affects community assembly can differ greatly for stable versus transient states. Differences arise because the contribution of such factors as mortality rate, environmental heterogeneity and plant-soil feedback to historical contingency changes as community assembly proceeds. We also show that transient states can last for a long time relative to immigration rate and generation time. These results argue for a conceptual shift of focus from alternative stable states to alternative transient states for understanding historical contingency in community assembly.     

Gradual regime shifts in spatially extended ecosystems

Ecosystem regime shifts are regarded as abrupt global transitions from one stable state to an alternative stable state, induced by slow environmental changes or by global disturbances. Spatially extended ecosystems, however, can also respond to local disturbances by the formation of small domains of the alternative state. Such a response can lead to gradual regime shifts involving front propagation and the coalescence of alternative-state domains. When one of the states is spatially patterned, a multitude of intermediate stable states appears, giving rise to step-like gradual shifts with extended pauses at these states. Using a minimal model, we study gradual state transitions and show that they precede abrupt transitions. We propose indicators to probe gradual regime shifts, and suggest that a combination of abrupt-shift indicators and gradual-shift indicators might be needed to unambiguously identify regime shifts. Our results are particularly relevant to desertification in drylands where transitions to bare soil take place from spotted vegetation, and the degradation process appears to involve step-like events of local vegetation mortality caused by repeated droughts.     

Coordinated approaches to quantify long‐term ecosystem dynamics in response to global change

Many serious ecosystem consequences of climate change will take decades or even centuries to emerge. Long‐term ecological responses to global change are strongly regulated by slow processes, such as changes in species composition, carbon dynamics in soil and by long‐lived plants, and accumulation of nutrient capitals. Understanding and predicting these processes require experiments on decadal time scales. But decadal experiments by themselves may not be adequate because many of the slow processes have characteristic time scales much longer than experiments can be maintained. This article promotes a coordinated approach that combines long‐term, large‐scale global change experiments with process studies and modeling. Long‐term global change manipulative experiments, especially in high‐priority ecosystems such as tropical forests and high‐latitude regions, are essential to maximize information gain concerning future states of the earth system. The long‐term experiments should be conducted in tandem with complementary process studies, such as those using model ecosystems, species replacements, laboratory incubations, isotope tracers, and greenhouse facilities. Models are essential to assimilate data from long‐term experiments and process studies together with information from long‐term observations, surveys, and space‐for‐time studies along environmental and biological gradients. Future research programs with coordinated long‐term experiments, process studies, and modeling have the potential to be the most effective strategy to gain the best information on long‐term ecosystem dynamics in response to global change.     

Coastal dynamics and wetlands stability. The Ebro delta case

Coastal wetland stability and structure can be significantly affected by littoral processes when they are close to the shoreline. Thus, under certain conditions, the combination of shoreline dynamics and direct wave action during storms can influence the stability of marshes and vegetation community composition. This interaction between littoral dynamics and coastal wetlands is illustrated by analysing processes taking place in the Buda Island (Ebro delta, NW Spain), where a coastal lagoon very close to a retreating shoreline exists. Two main time scales have been found to be relevant for interaction of coastal processes with ecosystem dynamics, the decadal and episodic scales. The decadal scale determines the average trend in beach width and directly controls the potential loss of wetland surface. The episodic scale is linked to the occurrence of wave and storm surge events and it determines a pulsing stress in the ecosystem through flooding, being key parameters to determine their direct influence the intensity and repetition of these events.