生态系统的多稳态与突变

许多生态系统可能在短时间内发生难以预料的状态突变, 其中一些生态系统突变的机理可以用多稳态理论进行解释。近年来生态系统的多稳态和突变现象及其机理吸引了研究者和管理者的广泛关注。本文重点对生态系统多稳态的理论基础、识别方法及稳态转换发生的早期预警信号进行综述, 并基于典型生态系统过程对现实世界中可能观测到的稳态转换进行实例分析, 最后对多稳态概念框架和理论应用中的潜在争议进行讨论, 以期为非线性生态系统动态的理论研究、管理实践和生物多样性保护等提供参考。     

Evidence of climate‐driven ecosystem reorganization in the Gulf of Mexico

The Gulf of Mexico is one of the most ecologically and economically valuable marine ecosystems in the world and is affected by a variety of natural and anthropogenic phenomena including climate, hurricanes, coastal development, agricultural runoff, oil spills, and fishing. These complex and interacting stressors, together with the highly dynamic nature of this ecosystem, present challenges for the effective management of its resources. We analyze a compilation of over 100 indicators representing physical, biological, and economic aspects of the Gulf of Mexico and find that an ecosystem‐wide reorganization occurred in the mid‐1990s. Further analysis of fishery landings composition data indicates a major shift in the late 1970s coincident with the advent of US national fisheries management policy, as well as significant shifts in the mid‐1960s and the mid‐1990s. These latter shifts are aligned temporally with changes in a major climate mode in the Atlantic Ocean: the Atlantic Multidecadal Oscillation (AMO). We provide an explanation for how the AMO may drive physical changes in the Gulf of Mexico, thus altering higher‐level ecosystem dynamics. The hypotheses presented here should provide focus for further targeted studies, particularly in regard to whether and how management should adjust to different climate regimes or states of nature. Our study highlights the challenges in understanding the effects of climatic drivers against a background of multiple anthropogenic pressures, particularly in a system where these forces interact in complex and nonlinear ways.     

Minor food sources can play a major role in secondary production in detritus‐based ecosystems

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What is the upper size limit for cosmopolitan distribution in free‐living microorganisms?

A comparison of testate amoebae assemblages from the Arctic and Antarctic (areas of similar habitat a maximum distance apart) is used to try and answer the question ‘What is the upper size limit for cosmopolitan distribution in free‐living microbes?’ Species restricted to either the Arctic or Antarctic exhibited sizes up to 230 μm while the largest cosmopolitan species was 135 μm in size. Comparison of the testate assemblages using a multivariate classificatory technique (TWINSPAN ) also suggested more restricted distribution for the larger species. There was a negative relationship between species size and number of sites at which it was recorded (r_s=?0.261, P < 0.05), with all the more widespread species having a size of below 100 μm. It is suggested that for testate amoebae cosmopolitan distributions become common below 100–150 μm. This suggests that most species of testate (indeed most free‐living microbes) have low species richness because of lack of opportunities for allopatric speciation as most are below 100 μm and so geographical isolation is unlikely. It is suggested that if this is correct, only the largest free‐living microbes (> 150 μm) are likely to be of conservation concern because of their smaller ranges. However, I point out that currently different studies are giving very different answers to the question, how ubiquitous and species rich are free‐living microbes? The subject requires further work to try and reconcile these different results.     

Habitat loss-induced tipping points in metapopulations with facilitation

Habitat loss is known to pervade extinction thresholds in metapopulations. Such thresholds result from a loss of stability that can eventually lead to collapse. Several models have been developed to understand the nature of these transitions and how they are affected by the locality of interactions, fluctuations or external drivers. Most models consider the impact of grazing or aridity as a control parameter that can trigger sudden shifts, once critical values are reached. Others explore instead the role played by habitat loss and fragmentation. Here we consider a minimal model incorporating facilitation between the individuals of the same species along with habitat destruction, with the aim of understanding how local cooperation and habitat loss interact with each other. A mathematical model incorporating facilitation and habitat destruction is derived, along with a spatially explicit simulation model. It is found that a catastrophic shift is expected for increasing levels of habitat loss, but the bifurcation becomes continuous when dispersal is local. Under these conditions, spatial patchiness is found and the qualitative change from discontinuous to continuous results are in agreement with previous studies on ecological systems. Our results suggest that species exhibiting facilitation and displaying short-range dispersal will be markedly more capable of avoiding catastrophic tipping points.     

Climate‐change impacts on sandy‐beach biota: crossing a line in the sand

Sandy ocean beaches are iconic assets that provide irreplaceable ecosystem services to society. Despite their great socioeconomic importance, beaches as ecosystems are severely under‐represented in the literature on climate‐change ecology. Here, we redress this imbalance by examining whether beach biota have been observed to respond to recent climate change in ways that are consistent with expectations under climate change. We base our assessments on evidence coming from case studies on beach invertebrates in South America and on sea turtles globally. Surprisingly, we find that observational evidence for climate‐change responses in beach biota is more convincing for invertebrates than for highly charismatic turtles. This asymmetry is paradoxical given the better theoretical understanding of the mechanisms by which turtles are likely to respond to changes in climate. Regardless of this disparity, knowledge of the unique attributes of beach systems can complement our detection of climate‐change impacts on sandy‐shore invertebrates to add rigor to studies of climate‐change ecology for sandy beaches. To this end, we combine theory from beach ecology and climate‐change ecology to put forward a suite of predictive hypotheses regarding climate impacts on beaches and to suggest ways that these can be tested. Addressing these hypotheses could significantly advance both beach and climate‐change ecology, thereby progressing understanding of how future climate change will impact coastal ecosystems more generally.     

Antarctic microbial diversity: the basis of polar ecosystem processes

Microorganisms are fundamental to the functioning of Antarctic ecosystems. Although microbial biomass can be immense in Southern Ocean blooms and freshwater cyanobacterial mats, species richness is generally more restricted than it is in temperate regions. However, there are representatives of a broad variety of taxa providing a diverse gene pool. Species diversity may be low while metabolic flexibility is high so that a few strains can provide most necessary functions. In this context, biodiversity is the sum of biological potential. This Special Issue highlights aspects of microbial ecology that can be studied only in Antarctica or which are defined most clearly in Antarctic habitats. Relatively simple microbial communities, or conspicuous species within them, can be used as indicators of microbial processes and responses to environmental change. These include the palaeological record of benthic diatoms and response of soil cyanobacterial communities to regional warming and UV-B stress. The climatic conditions and relict babitats of the Antarctic dry valleys are a valuable analogue for detecting microbial life and diversity on Mars. The global microbial biodiversity initiative Diversitas and international Antarctic networks such as BIOTAS (Biological Investigations of Terrestrial Antarctic Systems) harness taxonomic and ecophysiological expertize to understand better these unique polar ecosystems.     

Role of fungi in marine ecosystems

Marine fungi are an ecological rather than a taxonomic group and comprise an estimated 1500 species, excluding those that form lichens. They occur in most marine habitats and generally have a pantropical or pantemperate distribution. Marine fungi are major decomposers of woody and herbaceous substrates in marine ecosystems. Their importance lies in their ability to aggressively degrade lignocellulose. They may be important in the degradation of dead animals and animal parts. Marine fungi are important pathogens of plants and animals and also form symbiotic relationships with other organisms. The effect of disturbances on marine fungi is poorly investigated. Keystone marine species may exist, especially in mutualistic symbioses. However, as many saprophytes appear to carry out the same function simultaneously, they may be functionally redundant. The need for a concerted effort to investigate the biodiversity and role of marine fungi globally and on as many substrata as possible is presented.     

Global environmental changes: setting priorities for Latin American coastal habitats

As the effects of the Global Climate Changes on the costal regions of Central and South Americas advance, there is proportionally little research being made to understand such impacts. This commentary puts forward a series of propositions of strategies to improve performance of Central and South American science and policy making in order to cope with the future impacts of the Global Climate Changes in their coastal habitats.     

Consumer‐driven nutrient dynamics in freshwater ecosystems: from individuals to ecosystems

The role of animals in modulating nutrient cycling [hereafter, consumer‐driven nutrient dynamics (CND)] has been accepted as an important influence on both community structure and ecosystem function in aquatic systems. Yet there is great variability in the influence of CND across species and ecosystems, and the causes of this variation are not well understood. Here, we review and synthesize the mechanisms behind CND in fresh waters. We reviewed 131 articles on CND published between 1973 and 1 June 2015. The rate of new publications in CND has increased from 1.4 papers per year during 1973–2002 to 7.3 per year during 2003–2015. The majority of investigations are in North America with many concentrating on fish. More recent studies have focused on animal‐mediated nutrient excretion rates relative to nutrient demand and indirect impacts (e.g. decomposition). We identified several mechanisms that influence CND across levels of biological organization. Factors affecting the stoichiometric plasticity of consumers, including body size, feeding history and ontogeny, play an important role in determining the impact of individual consumers on nutrient dynamics and underlie the stoichiometry of CND across time and space. The abiotic characteristics of an ecosystem affect the net impact of consumers on ecosystem processes by influencing consumer metabolic processes (e.g. consumption and excretion/egestion rates), non‐CND supply of nutrients and ecosystem nutrient demand. Furthermore, the transformation and transport of elements by populations and communities of consumers also influences the flow of energy and nutrients across ecosystem boundaries. This review highlights that shifts in community composition or biomass of consumers and eco‐evolutionary underpinnings can have strong effects on the functional role of consumers in ecosystem processes, yet these are relatively unexplored aspects of CND. Future research should evaluate the value of using species traits and abiotic conditions to predict and understand the effects of consumers on ecosystem‐level nutrient dynamics across temporal and spatial scales. Moreover, new work in CND should strive to integrate knowledge from disparate fields of ecology and environmental science, such as physiology and ecosystem ecology, to develop a comprehensive and mechanistic understanding of the functional role of consumers. Comparative and experimental studies that develop testable hypotheses to challenge the current assumptions of CND, including consumer stoichiometric homeostasis, are needed to assess the significance of CND among species and across freshwater ecosystems.     

Global meta‐analysis of native and nonindigenous trophic traits in aquatic ecosystems

Ecologists have recently devoted their attention to the study of species traits and their role in the establishment and spread of nonindigenous species (NIS). However, research efforts have mostly focused on studies of terrestrial taxa, with lesser attention being dedicated to aquatic species. Aquatic habitats comprise of interconnected waterways, as well as exclusive introduction vectors that allow unparalleled artificial transport of species and their propagules. Consequently, species traits that commonly facilitate biological invasions in terrestrial systems may not be as represented in aquatic environments. We provide a global meta‐analysis of studies conducted in both marine and freshwater habitats. We selected studies that conducted experiments with native and NIS under common environmental conditions to allow detailed comparisons among species traits. In addition, we explored whether different factors such as species relatedness, functional feeding groups, latitude, climate, and experimental conditions could be linked to predictive traits. Our results show that species with traits that enhance consumption and growth have a substantially increased probability of establishing and spreading when entering novel ecosystems. Moreover, traits associated with predatory avoidance were more prevalent in NIS and therefore favour invasive species in aquatic habitats. When we analysed NIS interacting with taxonomically distinctive native taxa, we found that consumption and growth were particularly important traits. This suggests that particular attention should be paid to newly introduced species for which there are no close relatives in the local biota. Finally, we found a bias towards studies conducted in temperate regions, and thus, more studies in other climatic regions are needed. We conclude that studies aiming at predicting future range shifts should consider trophic traits of aquatic NIS as these traits are indicative of multiple interacting mechanisms involved in promoting species invasions.     

Role of fungi in freshwater ecosystems

There are more than 600 species of freshwater fungi with a greater number known from temperate, as compared to tropical, regions. Three main groups can be considered which include Ingoldian fungi, aquatic ascomycetes and non-Ingoldian hyphomycetes, chytrids and, oomycetes. The fungi occurring in lentic habitats mostly differ from those occurring in lotic habitats. Although there is no comprehensive work dealing with the biogeography of all groups of freshwater fungi, their distribution probably follows that of Ingoldian fungi, which are either cosmopolitan, restricted to pantemperate or pantropical regions, or in a few cases, have a restricted distribution. Freshwater fungi are thought to have evolved from terrestrial ancestors. Many species are clearly adapted to life in freshwater as their propagules have specialised aquatic dispersal abilities. Freshwater fungi are involved in the decay of wood and leafy material and also cause diseases of plants and animals. These areas are briefly reviewed. Gaps in our knowledge of freshwater fungi are discussed and areas in need of research are suggested.     

Tree diversity mitigates defoliation after a drought‐induced tipping point

Understanding the processes that underlie drought‐related tree vitality loss is essential for anticipating future forest dynamics, and for developing management plans aiming at increasing the resilience of forests to climate change. Forest vitality has been continuously monitored in Europe since the acid rain alert in the 1980s, and the intensive monitoring plots of ICP Forests offer the opportunity to investigate the effects of air pollution and climate change on forest condition. By making use of over 100 long‐term monitoring plots, where crown defoliation has been assessed extensively since 1990, we discovered a progressive shift from a negative to a positive effect of species richness on forest health. The observed tipping point in the balance of net interactions, from competition to facilitation, has never been reported from real ecosystems outside experimental conditions; and the strong temporal consistency of our observations with increasing drought stress emphasizes its climate change relevance. Furthermore, we show that higher species diversity has reduced the severity of defoliation in the long term. Our results confirm the greater resilience of diverse forests to future climate change‐induced stress. More generally, they add to an accumulating body of evidence on the large potential of tree species mixtures to face manifold disturbances in a changing world.     

Global change tipping points: above- and below-ground biotic interactions in a low diversity ecosystem

Low diversity ecosystems are expected to be more vulnerable to global changes although they have received less attention than high diversity ecosystems. Addressing the present state of the Antarctic Dry Valley region by focusing on the potential global changes that may alter the coupling of above- and below-ground species and ecosystem processes is a realistic and critical need that has value beyond the Antarctic community. Presented here are suggested implications of global change on the Dry Valley terrestrial systems and how these effects might be manifested in the future.     

Effects of climate‐driven primary production change on marine food webs: implications for fisheries and conservation

Climate change is altering the rate and distribution of primary production in the world's oceans. Primary production is critical to maintaining biodiversity and supporting fishery catches, but predicting the response of populations to primary production change is complicated by predation and competition interactions. We simulated the effects of change in primary production on diverse marine ecosystems across a wide latitudinal range in Australia using the marine food web model Ecosim. We link models of primary production of lower trophic levels (phytoplankton and benthic producers) under climate change with Ecosim to predict changes in fishery catch, fishery value, biomass of animals of conservation interest, and indicators of community composition. Under a plausible climate change scenario, primary production will increase around Australia and generally this benefits fisheries catch and value and leads to increased biomass of threatened marine animals such as turtles and sharks. However, community composition is not strongly affected. Sensitivity analyses indicate overall positive linear responses of functional groups to primary production change. Responses are robust to the ecosystem type and the complexity of the model used. However, model formulations with more complex predation and competition interactions can reverse the expected responses for some species, resulting in catch declines for some fished species and localized declines of turtle and marine mammal populations under primary productivity increases. We conclude that climate‐driven primary production change needs to be considered by marine ecosystem managers and more specifically, that production increases can simultaneously benefit fisheries and conservation. Greater focus on incorporating predation and competition interactions into models will significantly improve the ability to identify species and industries most at risk from climate change.     

Mammalian herbivores confer resilience of Arctic shrub‐dominated ecosystems to changing climate

Climate change is resulting in a rapid expansion of shrubs in the Arctic. This expansion has been shown to be reinforced by positive feedbacks, and it could thus set the ecosystem on a trajectory toward an alternate, more productive regime. Herbivores, on the other hand, are known to counteract the effects of simultaneous climate warming on shrub biomass. However, little is known about the impact of herbivores on resilience of these ecosystems, that is, the capacity of a system to absorb disturbance and still remain in the same regime, retaining the same function, structure, and feedbacks. Here, we investigated how herbivores affect resilience of shrub‐dominated systems to warming by studying the change of shrub biomass after a cessation of long‐term experimental warming in a forest–tundra ecotone. As predicted, warming increased the biomass of shrubs, and in the absence of herbivores, shrub biomass in tundra continued to increase 4 years after cessation of the artificial warming, indicating that positive effects of warming on plant growth may persist even over a subsequent colder period. Herbivores contributed to the resilience of these systems by returning them back to the original low‐biomass regime in both forest and tundra habitats. These results support the prediction that higher shrub biomass triggers positive feedbacks on soil processes and microclimate, which enable maintaining the rapid shrub growth even in colder climates. Furthermore, the results show that in our system, herbivores facilitate the resilience of shrub‐dominated ecosystems to climate warming.     

Greenhouse gas,upwelling‐favorable winds,and the future of coastal ocean upwelling ecosystems

Coastal ocean upwelling ecosystems generally represent the most productive large marine ecosystems of the world's oceans, in terms of both primary production rates and tonnages of exploitable fish produced. The Peruvian upwelling system, in particular, stands out as a major factor in world fish production. The Pacific trade winds have traditionally been considered to be the primary driving force for the upwelling system off Peru, but are projected to weaken as climate change proceeds. This leads to concern that the upwelling process in the Peru system, to which its productivity is linked, may likewise weaken. However, other mechanisms involving greenhouse‐associated intensification of thermal low‐pressure cells over the coastal landmasses of upwelling regions suggest general intensification of wind‐driven ocean upwelling in coastal upwelling regions of the world's oceans. But although certain empirical results have supported this expectation, it has not been consistently corroborated in climate model simulations, possibly because the scale of the coastal intensification may be small relative to the scales that are appropriately reflected in the standard models. Here we summarize available evidence for the intensification mechanism and present a proxy test that uses variations in water vapor, the dominant natural greenhouse gas, to offer multiple‐realization empirical evidence for action of the proposed mechanism in the real world situation. While many potential consequences to the future of marine ecosystems would codepend on climate change‐related changes in the thermocline and nutricline structures, an important subset, involving potential increased propensities for hypoxia, noxious gas eruptions, toxic red tide blooms, and/or jellyfish outbreaks, may depend more directly on changes in the upwelling‐favorable wind itself. A prospective role of fisheries in either mitigating or reinforcing this particular class of effects is suggested.     

Sea‐ice loss boosts visual search: fish foraging and changing pelagic interactions in polar oceans

Light is a central driver of biological processes and systems. Receding sea ice changes the lightscape of high‐latitude oceans and more light will penetrate into the sea. This affects bottom‐up control through primary productivity and top‐down control through vision‐based foraging. We model effects of sea‐ice shading on visual search to develop a mechanistic understanding of how climate‐driven sea‐ice retreat affects predator–prey interactions. We adapt a prey encounter model for ice‐covered waters, where prey‐detection performance of planktivorous fish depends on the light cycle. We use hindcast sea‐ice concentrations (past 35 years) and compare with a future no‐ice scenario to project visual range along two south–north transects with different sea‐ice distributions and seasonality, one through the Bering Sea and one through the Barents Sea. The transect approach captures the transition from sub‐Arctic to Arctic ecosystems and allows for comparison of latitudinal differences between longitudes. We find that past sea‐ice retreat has increased visual search at a rate of 2.7% to 4.2% per decade from the long‐term mean; and for high latitudes, we predict a 16‐fold increase in clearance rate. Top‐down control is therefore predicted to intensify. Ecological and evolutionary consequences for polar marine communities and energy flows would follow, possibly also as tipping points and regime shifts. We expect species distributions to track the receding ice‐edge, and in particular expect species with large migratory capacity to make foraging forays into high‐latitude oceans. However, the extreme seasonality in photoperiod of high‐latitude oceans may counteract such shifts and rather act as a zoogeographical filter limiting poleward range expansion. The provided mechanistic insights are relevant for pelagic ecosystems globally, including lakes where shifted distributions are seldom possible but where predator–prey consequences would be much related. As part of the discussion on photoperiodic implications for high‐latitude range shifts, we provide a short review of studies linking physical drivers to latitudinal extent.