Niche width collapse in a resilient top predator following ecosystem fragmentation

Much research has focused on identifying species that are susceptible to extinction following ecosystem fragmentation, yet even those species that persist in fragmented habitats may have fundamentally different ecological roles than conspecifics in unimpacted areas. Shifts in trophic role induced by fragmentation, especially of abundant top predators, could have transcendent impacts on food web architecture and stability, as well as ecosystem function. Here we use a novel measure of trophic niche width, based on stable isotope ratios, to assess effects of aquatic ecosystem fragmentation on trophic ecology of a resilient, dominant, top predator. We demonstrate collapse in trophic niche width of the predator in fragmented systems, a phenomenon related to significant reductions in diversity of potential prey taxa. Collapsed niche width reflects a homogenization of energy flow pathways to top predators, likely serving to destabilize remnant food webs and render apparently resilient top predators more susceptible to extinction through time.     

Modelling metapopulations with stochastic membrane systems

Metapopulations, or multi-patch systems, are models describing the interactions and the behavior of populations living in fragmented habitats. Dispersal, persistence and extinction are some of the characteristics of interest in ecological studies of metapopulations. In this paper, we propose a novel method to analyze metapopulations, which is based on a discrete and stochastic modelling framework in the area of Membrane Computing. New structural features of membrane systems, necessary to appropriately describe a multi-patch system, are introduced, such as the reduction of the maximal parallel consumption of objects, the spatial arrangement of membranes and the stochastic creation of objects. The role of the additional features, their meaning for a metapopulation model and the emergence of relevant behaviors are then investigated by means of stochastic simulations. Conclusive remarks and ideas for future research are finally presented.     

Demographic noise and resilience in a semi-arid ecosystem model

The scarcity of water characterising drylands forces vegetation to adopt appropriate survival strategies. Some of these generate water–vegetation feedback mechanisms that can lead to spatial self-organisation of vegetation, as it has been shown with models representing plants by a density of biomass, varying continuously in time and space. However, although plants are usually quite plastic they also display discrete qualities and stochastic behaviour. These features may give rise to demographic noise, which in certain cases can influence the qualitative dynamics of ecosystem models. In the present work we explore the effects of demographic noise on the resilience of a model semi-arid ecosystem. We introduce a spatial stochastic eco-hydrological hybrid model in which plants are modelled as discrete entities subject to stochastic dynamical rules, while the dynamics of surface and soil water are described by continuous variables. The model has a deterministic approximation very similar to previous continuous models of arid and semi-arid ecosystems. By means of numerical simulations we show that demographic noise can have important effects on the extinction and recovery dynamics of the system. In particular we find that the stochastic model escapes extinction under a wide range of conditions for which the corresponding deterministic approximation predicts absorption into desert states.     

Short-term transformation of matrix into hospitable habitat facilitates gene flow and mitigates fragmentation

Habitat fragmentation has major implications for demography and genetic structure of natural plant and animal populations as small and isolated populations are more prone to extinction. Therefore, many recent studies focus on spatial fragmentation. However, the temporal configuration of suitable habitat may also influence dispersal and gene flow in fragmented landscapes. We hypothesize that short-term switching of inhospitable matrix areas into suitable habitat can mitigate effects of spatial fragmentation in natural and seminatural ecosystems. To test our hypothesis, we investigated the hairy-footed gerbil (Gerbillurus paeba, Smith 1836), a ground-dwelling rodent, in fragmented Kalahari savannah areas. Here, rare events of high above mean annual rainfall suggest short-term matrix suitability. During the field survey in 'matrix' areas in the Kalahari (shrub encroachment by heavy grazing) we never observed the hairy-footed gerbil in years of average rainfall, but observed mass occurrences of this species during rare events of exceptionally high rainfall. In a second step, we developed an agent-based model simulating subpopulations in two neighbouring habitats and the separating matrix. Our mechanistic model reproduces the mass occurrences as observed in the field and thus suggests the possibly underlying processes. In particular, the temporary improvement in matrix quality allows reproduction in the matrix, thereby causing a substantial increase in population size. The model demonstrates further how the environmental trigger (rainfall) impacts genetic connectivity of two separated subpopulations. We identified seasonality as a driver of fragmentation but stochasticity leading to higher connectivity. We found that our concept of temporal fragmentation can be applied to numerous other fragmented populations in various ecological systems and provide examples from recent literature. We conclude that temporal aspects of fragmentation must be considered in both ecological research and conservation management.     

The influence of species diversity on ecosystem productivity: how,where, and why?

The effect of species diversity on ecosystem productivity is controversial, in large part because field experiments investigating this relationship have been fraught with difficulties. Unfortunately, there are few guidelines to aid researchers who must overcome these difficulties and determine whether global species losses seriously threaten the ecological and economic bases of terrestrial ecosystems. In response, I offer a set of hypotheses that describe how diversity might influence productivity in plant communities based on three well-known mechanisms: complementarity, facilitation, and the sampling effect. Emphasis on these mechanisms reveals the sensitivity of any diversity-productivity relationship to ecological context (i.e., where this relationship should be found); ecological context includes characteristics of the surrounding environment, temporal and spatial scales of observation, and the intensity of human management. In particular, the legitimacy of the sampling effect as a mechanism of productivity enhancement is dependent upon the degree to which stochastic events influence immigration and extinction processes in a given ecosystem. A mechanistic approach also requires that the three mechanisms be separated and quantified in diversity experiments, and I examine the most appropriate analyses for doing so, focusing on the overyielding technique. Finally, I question why productivity per se is a relevant management concern in non-agricultural systems once relationships among diversity, productivity, and the qualities of the surrounding environment are considered.     

Optimal control of population recovery – the role of economic restoration threshold

A variety of ecological systems around the world have been damaged in recent years, either by natural factors such as invasive species, storms and global change or by direct human activities such as overfishing and water pollution. Restoration of these systems to provide ecosystem services entails significant economic benefits. Thus, choosing how and when to restore in an optimal fashion is important, but has not been well studied. Here we examine a general model where population growth can be induced or accelerated by investing in active restoration. We show that the most cost‐effective method to restore an ecosystem dictates investment until the population approaches an ‘economic restoration threshold’, a density above which the ecosystem should be left to recover naturally. Therefore, determining this threshold is a key general approach for guiding efficient restoration management, and we demonstrate how to calculate this threshold for both deterministic and stochastic ecosystems.     

Disentangling the importance of ecological niches from stochastic processes across scales

Deterministic theories in community ecology suggest that local, niche-based processes, such as environmental filtering, biotic interactions and interspecific trade-offs largely determine patterns of species diversity and composition. In contrast, more stochastic theories emphasize the importance of chance colonization, random extinction and ecological drift. The schisms between deterministic and stochastic perspectives, which date back to the earliest days of ecology, continue to fuel contemporary debates (e.g. niches versus neutrality). As illustrated by the pioneering studies of Robert H. MacArthur and co-workers, resolution to these debates requires consideration of how the importance of local processes changes across scales. Here, we develop a framework for disentangling the relative importance of deterministic and stochastic processes in generating site-to-site variation in species composition (β-diversity) along ecological gradients (disturbance, productivity and biotic interactions) and among biogeographic regions that differ in the size of the regional species pool. We illustrate how to discern the importance of deterministic processes using null-model approaches that explicitly account for local and regional factors that inherently create stochastic turnover. By embracing processes across scales, we can build a more synthetic framework for understanding how niches structure patterns of biodiversity in the face of stochastic processes that emerge from local and biogeographic factors.     

Testing metapopulation dynamics using genetic, demographic and ecological data

The metapopulation concept is a cornerstone in the recent history of ecology and evolution. However, determining whether a natural system fits a metapopulation model is a complex issue. Extinction-colonization dynamics are indeed often difficult to quantify because species detectability is not always 100%, resulting in an imperfect record of extinctions. Here, we explore whether combining population genetics with demographic and ecological surveys can yield more realistic estimates of metapopulation dynamics. We apply this approach to the freshwater snail Drepanotrema depressissimum in a fragmented landscape of tropical ponds. In addition to studying correlations between genetic diversity and demographical or ecological characteristics, we undertake, for the first time, a detailed search for genetic signatures of extinction-recolonization events using temporal changes in allele frequencies within sites. Surprisingly, genetic data indicate that extinction is much rarer than suggested by demographic surveys. Consequently, this system is better described as a set of populations with different sizes and immigration rates than as a true metapopulation. We identify several cases of apparent extinction owing to nondetection of low-density populations, and of aestivating individuals in desiccated ponds. More generally, we observed a frequent mismatch between genetic and demographical/ecological information at small spatial and temporal scales. We discuss the causes of these discrepancies and show how these two types of data provide complementary information on population dynamics and history, especially when temporal genetic samples are available.     

Allee effects and resilience in stochastic populations

Allee effects, or positive functional relationships between a population’s density (or size) and its per unit abundance growth rate, are now considered to be a widespread if not common influence on the growth of ecological populations. Here we analyze how stochasticity and Allee effects combine to impact population persistence. We compare the deterministic and stochastic properties of four models: a logistic model (without Allee effects), and three versions of the original model of Allee effects proposed by Vito Volterra representing a weak Allee effect, a strong Allee effect, and a strong Allee effect with immigration. We employ the diffusion process approach for modeling single-species populations, and we focus on the properties of stationary distributions and of the mean first passage times. We show that stochasticity amplifies the risks arising from Allee effects, mainly by prolonging the amount of time a population spends at low abundance levels. Even weak Allee effects become consequential when the ubiquitous stochastic forces affecting natural populations are accounted for in population models. Although current concepts of ecological resilience are bound up in the properties of deterministic basins of attraction, a complete understanding of alternative stable states in ecological systems must include stochasticity.     

Bio-economic resource management under threats of environmental catastrophes

We combine ecological and economic dynamics to study the management of a natural resource that supports both ecosystem and human needs. Shrinking the resource base introduces a threat of occurrence of catastrophic ecological events, such as sudden ecosystem collapse. The occurrence conditions involve uncertainty of various types, and the distinction among these types is important for optimal resource management. When uncertainty is due to our ignorance of some aspects of the underlying ecology, the isolated equilibrium states characterizing optimal exploitation for many renewable resource problems become equilibrium intervals. Genuinely stochastic events shift the optimal equilibrium states, but maintain the structure of isolated equilibria.     

面向生态监管的多等级生态功能网格概念与框架

科学有效的生态监管是落实我国生态文明建设、保障区域生态安全、实现城市可持续发展的必要途径和重要抓手。生态系统的监管涉及到社会、经济、自然等多个维度,水、土、气、生等多类要素,国家到局地等多个尺度。目前,已有生态监管常存在生态系统、生态要素、行政单元的割裂。如何整体考虑以上特征,开展科学的、系统的、空间显性的生态监管亟需理论技术的创新。通过整合等级斑块动态范式、复合生态系统理论、多功能景观理论,构建了多等级生态功能网格框架,以期为生态监管提供一个综合的概念与技术框架。等级斑块动态范式从等级作用的角度构建了多等级网格骨架,复合生态系统理论从多个维度丰富了多等级网格内涵,多功能景观理论明确了网格的功能。通过耦合多等级生态功能网格的划分、评价和监管,可实现系统、整体、差异化的生态监管,为生态文明建设提供有力支撑。     

Disturbance-level-dependent post-disturbance succession in a Eurasian steppe

Anthropogenic disturbances may decrease as we take measures to control them. However, the patterns and mechanisms of post-disturbance ecosystem succession have rarely been studied. Here we reported that disturbance level determined the importance of stochastic relative to deterministic changes in ecosystem components (plant community composition, soil microbial community composition, and soil physicochemical indices), and thus predefined the pattern of post-disturbance ecosystem succession. We proposed a theoretical framework with five disturbance levels corresponding to distinct succession patterns. We conducted a nitrogen addition experiment in a temperate steppe, monitored these ecosystem components during “disturbance” treatment (2010–2014) and post-treatment “succession” (2014–2018). The disturbance level experienced by each component in each treatment was inferred by fitting the observed succession patterns into the theoretical framework. The mean disturbance level of these components was found to increase quadratically with nitrogen addition rate. This was because increasing nitrogen addition reduced the importance of stochastic relative to deterministic changes in these components, and these changes had a quadratic relationship with disturbance level. Overall, our results suggested that by monitoring the importance of stochastic relative to deterministic changes in an ecosystem, we can estimate disturbance levels and predict succession patterns, as well as propose disturbance-level-dependent strategies for post-disturbance restoration.

     

Modeling ecological traps for the control of feral pigs

Ecological traps are habitat sinks that are preferred by dispersing animals but have higher mortality or reduced fecundity compared to source habitats. Theory suggests that if mortality rates are sufficiently high, then ecological traps can result in extinction. An ecological trap may be created when pest animals are controlled in one area, but not in another area of equal habitat quality, and when there is density‐dependent immigration from the high‐density uncontrolled area to the low‐density controlled area. We used a logistic population model to explore how varying the proportion of habitat controlled, control mortality rate, and strength of density‐dependent immigration for feral pigs could affect the long‐term population abundance and time to extinction. Increasing control mortality, the proportion of habitat controlled and the strength of density‐dependent immigration decreased abundance both within and outside the area controlled. At higher levels of these parameters, extinction was achieved for feral pigs. We extended the analysis with a more complex stochastic, interactive model of feral pig dynamics in the Australian rangelands to examine how the same variables as the logistic model affected long‐term abundance in the controlled and uncontrolled area and time to extinction. Compared to the logistic model of feral pig dynamics, the stochastic interactive model predicted lower abundances and extinction at lower control mortalities and proportions of habitat controlled. To improve the realism of the stochastic interactive model, we substituted fixed mortality rates with a density‐dependent control mortality function, empirically derived from helicopter shooting exercises in Australia. Compared to the stochastic interactive model with fixed mortality rates, the model with the density‐dependent control mortality function did not predict as substantial decline in abundance in controlled or uncontrolled areas or extinction for any combination of variables. These models demonstrate that pest eradication is theoretically possible without the pest being controlled throughout its range because of density‐dependent immigration into the area controlled. The stronger the density‐dependent immigration, the better the overall control in controlled and uncontrolled habitat combined. However, the stronger the density‐dependent immigration, the poorer the control in the area controlled. For feral pigs, incorporating environmental stochasticity improves the prospects for eradication, but adding a realistic density‐dependent control function eliminates these prospects.     

Colonization and extinction dynamics of an annual plant metapopulation in an urban environment

The metapopulation framework considers that the spatiotemporal distribution of organisms results from a balance between the colonization and extinction of populations in a suitable and discrete habitat network. Recent spatially realistic metapopulation models have allowed patch dynamics to be investigated in natural populations but such models have rarely been applied to plants. Using a simple urban fragmented population system in which favourable habitat can be easily mapped, we studied patch dynamics in the annual plant Crepis sancta (Asteraceae). Using stochastic patch occupancy models (SPOMs) and multi‐year occupancy data we dissected extinction and colonization patterns in our system. Overall, our data were consistent with two distinct metapopulation scenarios. A metapopulation (sensu stricto) dynamic in which colonization occurs over a short distance and extinction is lowered by nearby occupied patches (rescue effect) was found in a set of patches close to the city centre, while a propagule rain model in which colonization occurs from a large external population was most consistent with data from other networks. Overall, the study highlights the importance of external seed sources in urban patch dynamics. Our analysis emphasizes the fact that plant distributions are governed not only by habitat properties but also by the intrinsic properties of colonization and dispersal of species. The metapopulation approach provides a valuable tool for understanding how colonization and extinction shape occupancy patterns in highly fragmented plant populations. Finally, this study points to the potential utility of more complex plant metapopulation models than traditionally used for analysing ecological and evolutionary processes in natural metapopulations.     

Advancing urban ecological studies: Frameworks,concepts, and results from the Baltimore Ecosystem Study

Abstract Urban ecological studies have had a long history, but they have not been a component of mainstream ecology until recently. The growing interest of ecologists in urban systems provides an opportunity to articulate integrative frameworks, and identify research tools and approaches that can help achieve a broader ecological understanding of urban systems. Based on our experience in the Baltimore Ecosystem Study (BES), Long‐term Ecological Research project, located in metropolitan Baltimore, Maryland, USA, we identify several frameworks that may be useful in comparative urban studies, and may be worthy of consideration in other integrative urban ecosystem studies: (i) spatial patch dynamics of biophysical and social factors; (ii) the watershed as an integrative tool; and (iii) the human ecosystem framework. These frameworks build on empirical research investigating urban biota, nutrient and energy budgets, ecological footprints of cities, as well as biotic classifications aimed at urban planning. These frameworks bring together perspectives, measurements, and models from biophysical and social sciences. We illustrate their application in the BES, which is designed to investigate (i) the structure and change of the urban ecosystem; (ii) the fluxes of matter, energy, capital, and population in the metropolis; and (iii) how ecological information affects the quality of the local and regional environments. Exemplary results concerning urban stream nutrient flux, the ability of riparian zones to process nitrate pollution, and the lags in the relationships between vegetation structure and socio‐economic factors in specific neighbourhoods are presented. The current advances in urban ecological studies have profited greatly from the variety of integrative frameworks and tools that have been tested and applied in urban areas over the last decade. The field is poised to make significant progress as a result of ongoing conceptual and empirical consolidation.     

Applying landscape metrics to species distribution model predictions to characterize internal range structure and associated changes

Distributional shifts in species ranges provide critical evidence of ecological responses to climate change. Assessments of climate-driven changes typically focus on broad-scale range shifts (e.g. poleward or upward), with ecological consequences at regional and local scales commonly overlooked. While these changes are informative for species presenting continuous geographic ranges, many species have discontinuous distributions—both natural (e.g. mountain or coastal species) or human-induced (e.g. species inhabiting fragmented landscapes)—where within-range changes can be significant. Here, we use an ecosystem engineer species (Sabellaria alveolata) with a naturally fragmented distribution as a case study to assess climate-driven changes in within-range occupancy across its entire global distribution. To this end, we applied landscape ecology metrics to outputs from species distribution modelling (SDM) in a novel unified framework. SDM predicted a 27.5% overall increase in the area of potentially suitable habitat under RCP 4.5 by 2050, which taken in isolation would have led to the classification of the species as a climate change winner. SDM further revealed that the latitudinal range is predicted to shrink because of decreased habitat suitability in the equatorward part of the range, not compensated by a poleward expansion. The use of landscape ecology metrics provided additional insights by identifying regions that are predicted to become increasingly fragmented in the future, potentially increasing extirpation risk by jeopardising metapopulation dynamics. This increased range fragmentation could have dramatic consequences for ecosystem structure and functioning. Importantly, the proposed framework—which brings together SDM and landscape metrics—can be widely used to study currently overlooked climate-driven changes in species internal range structure, without requiring detailed empirical knowledge of the modelled species. This approach represents an important advancement beyond predictive envelope approaches and could reveal itself as paramount for managers whose spatial scale of action usually ranges from local to regional.     

Natural extinction on islands

Almost all recent extinction of species or subspecies on islands comes from human activities. On the other hand, in local populations there is much natural extinction and immigration, i.e. turnover, on small islands. Most of this turnover occurs in locally rare species, and attests to the phenomenon of minimum viable population size. The MacArthur-Wilson theory is based on this turnover which, from an ecological point of view, is generally trivial. More useful theories of minimum viable population size are being developed. Rarity is the precursor of extinction, and species can be rare in several ways. Models of these phenomena are still primitive, particularly those that relate habitat availability to population density. Models of interactive communities show phenomena that may be relevant to the understanding of extinction in the geological record. Lotka-Volterra equations indicate considerable sensitivity to invasions, sometimes producing a cascade of extinction. Chemostat equations show that the behaviour of food chains can change dramatically with small changes in parameters, suggesting that small environmental effects can sometimes cause large ecological changes, including extinctions, in interactive biotic communities.     

集合生态系统研究15年回顾与展望

集合生态系统(meta-ecosystem)由法国的Loreau教授等于2003年提出,是指"由跨生态系统边界的物质流、能量流和生物体流所连接起来的一系列生态系统的集合",是对只关注生物体(organism)迁移交换的集合种群(meta-population)和集合群落(meta-community)概念的外推,也是为了给生态系统空间异质性研究提供一个重要的分析路径,对研究和理解生态系统的结构、过程、功能和异质性具有重要意义。通过对相关文献的梳理分析,简述了集合生态系统研究的基本状况,分析了对集合生态系统概念的狭义和广义两种理解,指出了探讨集合生态系统结构的两个方向,构建了分析集合生态系统研究的六维整体框架,综述了研究集合生态系统的两类方法,探讨了经验化的集合生态系统(empirical meta-ecosystem)的3种空间结构和两种构建路径。将集合生态系统概念和理论引入流域复合生态系统(integrated watershed ecosystem)的分析,为流域生态学研究提供新的概念框架。     

生境破碎化对动物种群存活的影响

生境破碎是生物多样性下降的主要原因之一。通常以岛屿生物地理学、异质种群生物学和景观生态学的理论来解释不同空间尺度中生境破碎化的生态学效应。生境破碎化引起面积效应、隔离效应和边缘效应。这些效应通过影响动物种群的绝灭阈值、分布和多度、种间关系以及生态系统过程,最终影响动物种群的存活。野外研究表明,破碎化对动物的影响,因物种、生境类型和地理区域不同而有所变化,因此,预测物种在破碎生境中的存活比较困难。研究热点集中于：确定生境面积损失和生境斑块的空间格局对破碎景观中物种绝灭的相对影响,破碎景观中物种的适宜生境比例和绝灭阈值,异质种群动态以及生态系统的生态过程。随着3S技术的发展,生境破碎化模型趋于复杂,而发展有效的模型和验证模型将成为一项富有挑战性的任务。     

Individual-based vs deterministic models for macroparasites: host cycles and extinction

Our understanding of the qualitative dynamics of host-macroparasite systems is mainly based on deterministic models. We study here an individual-based stochastic model that incorporates the same assumptions as the classical deterministic model. Stochastic simulations, using parameter values based on some case studies, preserve many features of the deterministic model, like the average value of the variables and the approximate length of the cycles.An important difference is that, even when deterministic models yield damped oscillations, stochastic simulations yield apparently sustained oscillations. The amplitude of such oscillations may be so large to threaten parasites' persistence.With density-dependence in parasite demographic traits, persistence increases somewhat. Allowing instead for infections from an external parasite reservoir, we found that host extinction may easily occur. However, the extinction probability is almost independent of the level of external infection over a wide intermediate parameter region.