Multiplicity of stable states in freshwater systems

It is shown with the use of minimal models that several ecological relationships in freshwater systems potentially give rise to the existence of alternative equilibria over a certain range of nutrient values. The existence of alternative stable states has some implications for the management of such systems. An important consequence is that signs of eutrophication are only apparent after the occurrence of changes that are very difficult to reverse. Reduction of the nutrient level as a measure to restore such systems gives poor results, but biomanipulation as an additional measure can have significant effects, provided that the nutrient level has been reduced enough to allow the existence of a stable alternative clear water equilibrium.     

Alternative stable states in communities with intraguild predation

Intraguild predation (IGP) is a widespread ecological phenomenon in which two consumers that share a resource also engage in a predator-prey interaction. Theory on IGP predicts the occurrence of alternative stable states, but empirical evidence of such states is scarce. This raises the question of whether alternative states are a rare phenomenon that is unlikely to be observed in nature. Here we analyze a model in which the resource exhibits logistic or chemostat dynamics and consumers have saturating (Type II) functional responses. We show that alternative states can arise for a wide range of biological scenarios and that environmental constraints can make their detection difficult. Our analysis identifies three possible combinations of alternative states: (i) IG prey or IG predator, (ii) coexistence or IG predator, and (iii) coexistence or IG prey. Bifurcation diagrams reveal that alternative states are possible over large regions of the parameter space. However, they can be limited to narrow ranges along the resource productivity axis, which may make it difficult to observe the occurrence of alternative states in communities with IGP. Microcosm experiments provide a promising avenue for detecting combinations of asymptotically stable states along a productivity gradient.     

Tracking unstable states: ecosystem dynamics in a changing world

Ecological systems can show complex and sometimes abrupt responses to environmental change, with important implications for their resilience. Theories of alternate stable states have been used to predict regime shifts of ecosystems as equilibrium responses to sufficiently slow environmental change. The actual rate of environmental change is a key factor affecting the response, yet we are still lacking a non-equilibrium theory that explicitly considers the influence of this rate of environmental change. We present a metacommunity model of predator–prey interactions displaying multiple stable states, and we impose an explicit rate of environmental change in habitat quality (carrying capacity) and connectivity (dispersal rate). We study how regime shifts depend on the rate of environmental change and compare the outcome with a stability analysis in the corresponding constant environment. Our results reveal that in a changing environment, the community can track states that are unstable in the constant environment. This tracking can lead to regime shifts, including local extinctions, that are not predicted by alternative stable state theory. In our metacommunity, tracking unstable states also controls the maintenance of spatial heterogeneity and spatial synchrony. Tracking unstable states can also lead to regime shifts that may be reversible or irreversible. Our study extends current regime shift theories to integrate rate-dependent responses to environmental change. It reveals the key role of unstable states for predicting transient dynamics and long-term resilience of ecological systems to climate change.     

Detection of alternative stable states in marine communities

During the last 20 years there have been several attempts to test the theory of alternative stable states using marine systems. The results have been mixed, and there have been conflicting interpretations of the outcomes. In an attempt to resolve some of the problems, the theory and evidence for alternative stable states are reviewed. There are several different views of what is meant by alternative stable states, and so the key elements of the theory of alternative stable states are set in the context of marine systems. Appropriate experimental designs for detecting alternative states are discussed, and recent experimental studies in coral reefs, rocky intertidal systems, soft-sediment beds, and subtidal systems are briefly reviewed. Suggestions for improving experimental designs are proposed and unresolved issues are highlighted.     

Regime shifts in exploited marine food webs: detecting mechanisms underlying alternative stable states using size-structured community dynamics theory

Many marine ecosystems have undergone ‘regime shifts’, i.e. abrupt reorganizations across trophic levels. Establishing whether these constitute shifts between alternative stable states is of key importance for the prospects of ecosystem recovery and for management. We show how mechanisms underlying alternative stable states caused by predator–prey interactions can be revealed in field data, using analyses guided by theory on size-structured community dynamics. This is done by combining data on individual performance (such as growth and fecundity) with information on population size and prey availability. We use Atlantic cod (Gadus morhua) and their prey in the Baltic Sea as an example to discuss and distinguish two types of mechanisms, ‘cultivation-depensation’ and ‘overcompensation’, that can cause alternative stable states preventing the recovery of overexploited piscivorous fish populations. Importantly, the type of mechanism can be inferred already from changes in the predators'' body growth in different life stages. Our approach can thus be readily applied to monitored stocks of piscivorous fish species, for which this information often can be assembled. Using this tool can help resolve the causes of catastrophic collapses in marine predatory–prey systems and guide fisheries managers on how to successfully restore collapsed piscivorous fish stocks.     

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.     

Seeking alternative stable states in a deep lake

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Alternative states in the phytoplankton of Lake Kinneret, Israel (Sea of Galilee)

1. Through analyses of a 34‐year record of phytoplankton, zooplankton and physicochemical parameters from Lake Kinneret, Israel, we show that distinct and persistent phytoplankton assemblage states occurred from winter to summer. 2. The most obvious characteristic of these states was the presence or absence of a spring bloom of the dinoflagellate, Peridinium gatunense. 3. Analyses of the data within the framework of the alternative states model revealed a possible complex triggering mechanism, and system hysteresis. 4. A change in zooplankton biomass and body size coincident with changes in predation pressure associated with the collapse of the Kinneret Bleak, Acanthobrama terraesanctae, fishery appeared to be the ‘slow changing’ variable in the context of the alternative states model. Alternative phytoplankton states were only possible after this variable crossed a threshold in 1993–94, following the collapse of the fishery. 5. When alternative states were possible, some physicochemical parameters and the structure of the zooplankton assemblage appeared to control which phytoplankton state emerged in a given year. In years without a P. gatunense bloom, important physicochemical parameters in winter included low NO₃ loading, high water temperature, high water level, a deeper thermocline, low transparency, high concentrations of NO₃ and Cl in the epilimnion, and low concentration of epilimnetic total phosphorus. In addition, the cladoceran Chydorus sphaericus and adults of the copepod Mesocyclops ogunnus were observed in winter in years without a bloom. 6. Zooplankton biomass and body size of some taxa have recovered since the 1993–94 collapse of the fishery, yet incidence of both phytoplankton states in Lake Kinneret was still possible. Within the framework of the alternative states model, this suggests that the slow changing variable threshold where alternative states became possible is different from the threshold where alternative states will no longer be possible. In other words, the system is characterised by a hysteresis.     

Feedbacks between phosphorus deposition and canopy cover: The emergence of multiple stable states in tropical dry forests

Dry forests represent a large percentage of tropical forests and are vulnerable to both anthropogenic and natural disturbances, yet important aspects of their sensitivity to disruption remain poorly understood. It is particularly unclear how changes in land-use or tropical storm patterns may affect the resiliency of phosphorus (P)-limited neotropical forests. In these systems, vegetation is sustained in the long-term by atmospheric P-inputs through rainfall, dust, or fog. Past research supports the idea that dust and fog deposition are dependent on canopy density (e.g. leaf area index). Thus, the canopy may function as a 'trap' for P, enabling a positive feedback between vegetation and P-deposition. We developed a conceptual model to investigate how Neotropical vegetation may respond to reduced P-deposition due to canopy losses. The model suggests that a canopy-deposition feedback may induce bistable vegetation dynamics; under some conditions, forests may be unable to naturally recover from relatively small disturbances.     

Alternative stable states in a wet calcareous dune slack in The Netherlands

Abstract. Evidence is presented for the occurrence of alternative stable states in a wet calcareous dune slack on the Frisian island of Texel, The Netherlands. An early pioneer stage (0.5 kgm^?2 total standing crop) and a more productive later successional stage (2.9 kg m^?2) occur side by side, with sharp boundaries between them. The pioneer vegetation has been recorded at the site for more than 62 yr. These features indicate the occurrence of a positive‐feedback mechanism that has led to alternative stable states. Analyses of ground and surface water composition, and decalcification depths, indicated that hydrologically the study site can be characterized as a flow‐through slack, with exfiltration of calcareous groundwater on one side and infiltration of surface water on the other side of the slack. These differences in hydrological conditions have led to distinct differences in environmental conditions within the dune slack. The occurrence of the two successional stages can, however, not be explained by differences in hydrological conditions since both stages occur side by side in the centre of the dune slack. It is, therefore, more likely that biotic interactions are the cause of the vegetation pattern. Three possible mechanisms for feedback processes are discussed: (1) enhanced nitrogen loss; (2) sulfide toxicity and (3) nutrient accumulation in internal cycle.     

Submerged macrophytes, zooplankton and the predominance of low- over high-chlorophyll states in western boreal, shallow-water wetlands

1. Shallow‐water (<2 m deep) wetlands in northern Alberta's western boreal forest tend to have either extremely high‐chlorophyll, pea‐green water or water that is low in chlorophyll and clear. The relative importance of top‐down processes (e.g. zooplankton grazing) compared with bottom‐up processes (e.g. nutrient limitation) for regulating the existence of these alternate states has yet to be explicitly investigated in these poorly studied waterbodies. 2. To assess the relationship between chemical and biological factors and the predominance of low‐chlorophyll over high‐chlorophyll states, a survey of 24 shallow‐water wetlands in northern Alberta was conducted over the summers of 2001 and 2002. 3. In wetlands without fish, statistical analysis indicated that high‐ and low‐chlorophyll sites could be differentiated with 82% accuracy based solely on whether submerged aquatic vegetation (SAV) coverage was greater or less than 25%. Further, high‐SAV lakes with zooplankton communities dominated by large cladocerans were clear 96% of the time. 4. In the few wetlands that supported stickleback populations, large cladocerans were absent. However, the development of zooplankton communities dominated by small cladocera (Bosmina) with calanoid copepods in fish‐containing wetlands corresponded with a shift from high‐ to low‐chlorophyll states. 5. Results suggest that in fishless wetlands high‐SAV coverage and grazing by large, SAV‐associated cladocerans promotes low‐chlorophyll states. Results also suggest that suppression of macroinvertebrate predators by sticklebacks allowing Bosmina to flourish may also promote low‐chlorophyll, clear‐water conditions.     

Positive feedback and alternative stable states in inbreeding, cooperation, sex roles and other evolutionary processes

A large proportion of studies in systems science focus on processes involving a mixture of positive and negative feedbacks, which are also common themes in evolutionary ecology. Examples of negative feedback are density dependence (population regulation) and frequency-dependent selection (polymorphisms). Positive feedback, in turn, plays a role in Fisherian 'runaway' sexual selection, the evolution of cooperation, selfing and inbreeding tolerance under purging of deleterious alleles, and the evolution of sex differences in parental care. All these examples feature self-reinforcing processes where the increase in the value of a trait selects for further increases, sometimes via a coevolutionary feedback loop with another trait. Positive feedback often leads to alternative stable states (evolutionary endpoints), making the interpretation of evolutionary predictions challenging. Here, we discuss conceptual issues such as the relationship between self-reinforcing selection and disruptive selection. We also present an extension of a previous model on parental care, focusing on the relationship between the operational sex ratio and sexual selection, and the influence of this relationship on the evolution of biparental or uniparental care.     

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.     

生态系统多稳态研究进展

在多稳态的生态系统中,外力可能导致生态系统状态突然之间发生不可逆转的转变,从而达到一个新的平衡状态。但目前对多稳态理论的系统研究很少,如何使用预警信号来预测生态系统的状态转变依旧是个难题。通过多稳态理论的梳理提出了一个更加综合的多稳态定义,并以放牧模型为例,系统总结了多稳态理论的相关概念,将多稳态理论应用在生态系统演替和扰沌理论的解释中;通过对生态系统稳态转换预警信号的原理、优缺点和应用条件的分析,对不同尺度下多稳态的研究方法进行了归纳;最后提出了目前多稳态领域的研究问题和未来的研究重点。结果表明:(1)将时间和空间预警信号结合在一起,并量化正确预警信号的概率,对错误预警信号的比例进行加权,可能会提供更准确的稳态转换的预报。(2)定量观测试验适用于小尺度的研究,而较大尺度的研究则采用简化的模型来模拟研究,选择正确的尺度极有可能改变预警信号的可靠性。(3)结合多稳态理论研究生态系统临界转换和反馈控制机制,并将基于性状的特征指标和进化动力学纳入其中,是生态系统修复实践的重要研究方向。(4)将多稳态相关理论和生态保护管理政策的实践相结合,是多稳态理论未来应用的前景。本研究为多稳态理论和实践的...     

Alternative stable states in eutrophic,shallow freshwater systems: A minimal model

A simple mathematical model was constructed, describing the relationships between pike, bream, aquatic macrophytes and the nutrient loading of shallow lakes. The model is analyzed with the use of zero-isoclines. It is concluded that, over a certain range of nutrient concentrations, the ecological relations incorporated in the model can give rise to the existence of two alternative stable equilibria;viz. a turbid bream-dominated one, and a clear state in which pike and aquatic vegetation are abundant. Under oligotrophic conditions, the clear-water state represents the only stable equilibrium; however, at high nutrient levels, the clear state is absent, or only locally stable. The response of the model to both increase and decrease of the nutrient level is characterised by hysteresis. The results indicate that manipulation of fish densities as a measure to improve water quality is only likely to produce long-term results when the nutrient level is below a certain threshold.     

Alternative stable equilibria and critical thresholds created by fire regimes and plant responses in a fire‐prone community

Wildfire is a dominant disturbance in many ecosystems, and fire frequency and intensity are being altered as climates change. Through effects on mortality and regeneration, fire affects plant community composition, species richness, and carbon cycling. In some regions, changes to fire regimes could result in critical, non‐reversible transitions from forest to non‐forested states. For example, the Klamath ecoregion (northwest United States) supports extensive conifer forests that are initially replaced by hardwood chaparral following high‐severity fire, but eventually return to conifer forest during the fire‐free periods. Climate change alters both the fire regime and post‐fire recovery dynamics, potentially causing shrubland to persist as a stable (i.e. self‐renewing) vegetation stage, rather than an ephemeral stage. Here, we present a theoretical investigation of how changes in plant traits and fire regimes can alter the stability of communities in forest‐shrub systems such as the Klamath. Our model captures the key characteristics of the system, including life‐stage‐specific responses to disturbance and asymmetrical competitive interactions. We assess vegetation stability via invasion analysis, and conclude that portions of the landscape that are currently forested also can be stable as shrubland. We identify parameter thresholds where community equilibria change from stable to unstable, and show how these thresholds may shift in response to changes in life‐history or environmental parameters. For instance, conifer maturation rates are expected to decrease as aridity increases under climate change, and our model shows that this reduction decreases the fire frequencies at which forests become unstable. Increases in fire activity sufficient to destabilize forest communities are likely to occur in more arid future climates. If widespread, this would result in reduced carbon stocks and a positive feedback to climate change. Changes in stability may be altered by management practices.