扩散对破碎化景观上宿主-寄生种群动态的影响

景观破碎化和扩散是空间种群模型的重要因素,对生物入侵存在着深远的影响。本章将基于偶对近似模型,探讨由局部和全局宿主-寄生相互作用共同决定的扩散模式对破坏性景观上疾病入侵与传播的影响。其中,生境破坏由生境丧失量与生境破碎化程度来描述。模拟结果显示,宿主和病毒的全局扩散对疾病的入侵与种群密度产生不对称效应：病毒的全局扩散对系统产生的影响较宿主的全局扩散更为显著。不同扩散模式下,生境丧失越高或破碎化程度越低,均将越有害于寄生病毒的入侵;同时,生境的破坏程度也显著地影响了入侵阈值对扩散模式的响应机制。本文研究结果暗示,景观破碎化的空间分布格局以及病毒扩散的限制均可作为物种保护与管理中有效的疾病控制策略。该研究结果在一定意义上丰富和发展了寄生感染理论,为物种保护提供了生态学理论依据。     

Non-neutral community dynamics: empirical predictions for ecosystem function and diversity from linearized consumer–resource interactions

A general model of linearized species interactions, essentially Lotka–Volterra theory, applied to questions of biodiversity has previously been shown to be a powerful tool for understanding local species–abundance patterns and community responses to environmental change for a single trophic level. Here this approach is extended to predict community composition and responses to environmental changes in trophically structured systems. We show how resource and consumer species richness and their relative abundances vary with the means and variances in enrichment level and strengths of intra- and interspecific interactions. Also demonstrated are the responses of local resource and consumer species richness to the global species pools at both trophic levels, as well as the covariation with net resource productivity. These predictions for resource and consumer specific responses to changes in environmental enrichment and global biodiversity are directly testable.     

Evolution of dispersal in a multi‐trophic community context

Much is known about the evolution of dispersal when species interact with their resources or natural enemies, but very little is known about dispersal evolution when species interact with both resources and natural enemies. Here I investigate how the dispersal of an intermediate consumer evolves in response to its interactions with a basal resource and top predator. I find that dispersal evolution is possible even when the consumer species is not directly affected by environmental variability, but rather experiences the consequences that such variability has on its resource and predator. Spatial variation in the consumer's fitness is driven by spatial heterogeneity in resource productivity, which determines whether a predator can colonize a resource‐consumer community. Temporal variation in the consumer's fitness is driven by random disturbances that cause periodic local extinctions of the predator, followed by recolonizations that lead to transient fluctuations in consumer abundance. When spatial variation in resource productivity is low and the predator can colonize all patches in the landscape, there is no spatial variation in consumer fitness but temporal variation in fitness favors the evolution of a dispersal monomorphism. When spatial variation in resource productivity is high and the predator cannot colonize many patches in the landscape, spatial variation in fitness selects against dispersal. In this case, temporal variation can promote the evolution of a dispersal polymorphism with sedentary and mobile phenotypes, but only for certain types of tri‐trophic interactions. This finding underscores the importance of indirect interactions in shaping the evolution of dispersal. While the ecological community can provide a strong selective environment for the evolution of dispersal, the nature of interactions between trophic levels can also impose constraints on evolution.     

Synchronization in ecological systems by weak dispersal coupling with time delay

One of the most salient spatiotemporal patterns in population ecology is the synchronization of fluctuating local populations across vast spatial extent. Synchronization of abundance has been widely observed across a range of spatial scales in relation to the rate of dispersal among discrete populations. However, the dependence of synchrony on patterns of among-patch movement across heterogeneous landscapes has been largely ignored. Here, we consider the duration of movement between two predator–prey communities connected by weak dispersal and its effect on population synchrony. More specifically, we introduce time-delayed dispersal to incorporate the finite transmission time between discrete populations across a continuous landscape. Reducing the system to a phase model using weakly connected network theory, it is found that the time delay is an important factor determining the nature and stability of phase-locked states. Our analysis predicts enhanced convergence to stable synchronous fluctuations in general and a decreased ability of systems to produce in-phase synchronization dynamics in the presence of delayed dispersal. These results introduce delayed dispersal as a tool for understanding the importance of dispersal time across a landscape matrix in affecting metacommunity dynamics. They further highlight the importance of landscape and dispersal patterns for predicting the onset of synchrony between weakly coupled populations.     

局域种群的Allee效应和集合种群的同步性

从包含Allee效应的局域种群出发,建立了耦合映像格子模型,即集合种群模型.通过分析和计算机模拟表明:(1)当局域种群受到Allee效应强度较大时,集合种群同步灭绝;(2)而当Allee效应强度相对较弱时,通过稳定局域种群动态(减少混沌)使得集合种群发生同步波动,而这种同步波动能够增加集合种群的灭绝风险;(3)斑块间的连接程度对集合种群同步波动的发生有很大的影响,适当的破碎化有利于集合种群的续存.全局迁移和Allee效应结合起来增加了集合种群同步波动的可能,从而增加集合种群的灭绝风险.这些结果对理解同步性的机理、利用同步机理来制定物种保护策略和害虫防治都有重要的意义.     

When can refuges mediate the genetic effects of fire regimes? A simulation study of the effects of topography and weather on neutral and adaptive genetic diversity in fire‐prone landscapes

Understanding how landscape heterogeneity mediates the effects of fire on biodiversity is increasingly important under global changes in fire regimes. We used a simulation experiment to investigate how fire regimes interact with topography and weather to shape neutral and selection‐driven genetic diversity under alternative dispersal scenarios, and to explore the conditions under which microrefuges can maintain genetic diversity of populations exposed to recurrent fire. Spatial heterogeneity in simulated fire frequency occurred in topographically complex landscapes, with fire refuges and fire‐prone “hotspots” apparent. Interannual weather variability reduced the effect of topography on fire patterns, with refuges less apparent under high weather variability. Neutral genetic diversity was correlated with long‐term fire frequency under spatially heterogeneous fire regimes, being higher in fire refuges than fire‐prone areas, except under high dispersal or low fire severity (low mortality). This generated different spatial genetic structures in fire‐prone and fire‐refuge components of the landscape, despite similar dispersal. In contrast, genetic diversity was only associated with time since the most recent fire in flat landscapes without predictable refuges and hotspots. Genetic effects of selection driven by fire‐related conditions depended on selection pressure, migration distance and spatial heterogeneity in fire regimes. Allele frequencies at a locus conferring higher fitness under successional environmental conditions followed a pattern of “temporal adaptation” to contemporary conditions under strong selection pressure and high migration. However, selected allele frequencies were correlated with spatial variation in long‐term mean fire frequency (relating to environmental predictability) under weak dispersal, low selection pressure and strong spatial heterogeneity in fire regimes.     

Climate change in metacommunities: dispersal gives double-sided effects on persistence

Climate change is increasingly affecting the structure and dynamics of ecological communities both at local and at regional scales, and this can be expected to have important consequences for their robustness and long-term persistence. The aim of the present work is to analyse how the spatial structure of the landscape and dispersal patterns of species (dispersal rate and average dispersal distance) affects metacommunity response to two disturbances: (i) increased mortality during dispersal and (ii) local species extinction. We analyse the disturbances both in isolation and in combination. Using a spatially and dynamically explicit metacommunity model, we find that the effect of dispersal on metacommunity persistence is two-sided: on the one hand, high dispersal significantly reduces the risk of bottom-up extinction cascades following the local removal of a species; on the other hand, when dispersal imposes a risk to the dispersing individuals, high dispersal increases extinction risks, especially when dispersal is global. Large-bodied species with long generation times at the highest trophic level are particularly vulnerable to extinction when dispersal involves a risk. This suggests that decreasing the mortality risk of dispersing individuals by improving the quality of the habitat matrix may greatly increase the robustness of metacommunities.     

Predicting food‐web structure with metacommunity models

Synthesis Metacommunity theory aims to elucidate the relative influence of local and regional‐scale processes in generating diversity patterns across the landscape. Metacommunity research has focused largely on assemblages of competing organisms within a single trophic level. Here, we test the ability of metacommunity models to predict the network structure of the aquatic food web found in the leaves of the northern pitcher plant Sarracenia purpurea. The species‐sorting and patch‐dynamics models most accurately reproduced nine food web properties, suggesting that local‐scale interactions play an important role in structuring Sarracenia food webs. Our approach can be applied to any well‐resolved food web for which data are available from multiple locations. The metacommunity framework explores the relative influence of local and regional‐scale processes in generating diversity patterns across the landscape. Metacommunity models and empirical studies have focused mostly on assemblages of competing organisms within a single trophic level. Studies of multi‐trophic metacommunities are predominantly restricted to simplified trophic motifs and rarely consider entire food webs. We tested the ability of the patch‐dynamics, species‐sorting, mass‐effects, and neutral metacommunity models, as well as three hybrid models, to reproduce empirical patterns of food web structure and composition in the complex aquatic food web found in the northern pitcher plant Sarracenia purpurea. We used empirical data to determine regional species pools and estimate dispersal probabilities, simulated local food‐web dynamics, dispersed species from regional pools into local food webs at rates based on the assumptions of each metacommunity model, and tested their relative fits to empirical data on food‐web structure. The species‐sorting and patch‐dynamics models most accurately reproduced nine food web properties, suggesting that local‐scale interactions were important in structuring Sarracenia food webs. However, differences in dispersal abilities were also important in models that accurately reproduced empirical food web properties. Although the models were tested using pitcher‐plant food webs, the approach we have developed can be applied to any well‐resolved food web for which data are available from multiple locations.     

Adapt or disperse: understanding species persistence in a changing world

The majority of studies on environmental change focus on the response of single species and neglect fundamental biotic interactions, such as mutualism, competition, predation, and parasitism, which complicate patterns of species persistence. Under global warming, disruption of community interactions can arise when species differ in their sensitivity to rising temperature, leading to mismatched phenologies and/or dispersal patterns. To study species persistence under global climate change, it is critical to consider the ecology and evolution of multispecies interactions; however, the sheer number of potential interactions makes a full study of all interactions unfeasible. One mechanistic approach to solving the problem of complicated community context to global change is to (i) define strategy groups of species based on life‐history traits, trophic position, or location in the ecosystem, (ii) identify species involved in key interactions within these groups, and (iii) determine from the interactions of these key species which traits to study in order to understand the response to global warming. We review the importance of multispecies interactions looking at two trait categories: thermal sensitivity of metabolic rate and associated life‐history traits and dispersal traits of species. A survey of published literature shows pronounced and consistent differences among trophic groups in thermal sensitivity of life‐history traits and in dispersal distances. Our approach increases the feasibility of unraveling such a large and diverse set of community interactions, with the ultimate goal of improving our understanding of community responses to global warming.     

Nutrient flows between ecosystems can destabilize simple food chains

Dispersal of organisms has large effects on the dynamics and stability of populations and communities. However, current metacommunity theory largely ignores how the flows of limiting nutrients across ecosystems can influence communities. We studied a meta-ecosystem model where two autotroph-consumer communities are spatially coupled through the diffusion of the limiting nutrient. We analyzed regional and local stability, as well as spatial and temporal synchrony to elucidate the impacts of nutrient recycling and diffusion on trophic dynamics. We show that nutrient diffusion is capable of inducing asynchronous local destabilization of biotic compartments through a diffusion-induced spatiotemporal bifurcation. Nutrient recycling interacts with nutrient diffusion and influences the susceptibility of the meta-ecosystem to diffusion-induced instabilities. This interaction between nutrient recycling and transport is further shown to depend on ecosystem enrichment. It more generally emphasizes the importance of meta-ecosystem theory for predicting species persistence and distribution in managed ecosystems.     

Thermal acclimation modulates the impacts of temperature and enrichment on trophic interaction strengths and population dynamics

Global change affects individual phenotypes and biotic interactions, which can have cascading effects up to the ecosystem level. However, the role of environmentally induced phenotypic plasticity in species interactions is poorly understood, leaving a substantial gap in our knowledge of the impacts of global change on ecosystems. Using a cladoceran–dragonfly system, we experimentally investigated the effects of thermal acclimation, acute temperature change and enrichment on predator functional response and metabolic rate. Using our experimental data, we next parameterized a population dynamics model to determine the consequences of these effects on trophic interaction strength and food‐chain stability. We found that (1) predation and metabolic rates of the dragonfly larvae increase with acute warming, (2) warm‐acclimated larvae have a higher maximum predation rate than cold‐acclimated ones, and (3) long‐term interaction strength increases with enrichment but decreases with both acclimation and acute temperatures. Overall, our experimental results show that thermal acclimation can buffer negative impacts of environmental change on predators and increase food‐web stability and persistence. We conclude that the effect of acclimation and, more generally, phenotypic plasticity on trophic interactions should not be overlooked if we aim to understand the effects of climate change and enrichment on species interaction strength and food‐web stability.     

Genetic structuring in fluctuating populations

The effect of genetic drift in spatially distributed dispersal-linked and density-regulated populations is studied in a classical one-locus two-allele system. We analyse emergence of genetic differentiation assuming random drift only, where the noise-like variability is due to demographic stochasticity. We find emergence of clusters of sub-units with local allele fixation and persistence of both alleles in lengthy simulations. We demonstrate that local allele fixation (extending over a number of adjoining spatial sub-units) – without global loss of alleles – may occur when the carrying capacities of local patches are small, under a full range population dynamic regimes, when dispersal rate is small, and when redistribution (through dispersal) does not act as global mixer. These results are novel. The key to the observations is that drift is simultaneously influenced by distance-dependent dispersal, demographic stochasticity and autocorrelated population fluctuations due to delayed-density dependence. These are standard elements of contemporary population models in spatially structured context. With stable large populations, no stochasticity and dispersal limited to neighbours only, our model collapses to the stepping-stone model, while with dispersal being random and global, the model collapses to Wright's island model.     

Use of stable carbon and nitrogen isotopes in insect trophic ecology

Insects are the most diverse organisms and often the most abundant animals in some ecosystems. Despite the importance of their functional roles and of the knowledge for conservation, the trophic ecology of many insect species is not fully understood. In this review, I present how stable carbon (C) and nitrogen (N) isotopes have been used to reveal the trophic ecology of insects over the last 30 years. The isotopic studies on insects have used differences in C isotope ratios between C₃ and C₄ plants, along vertical profiles of temperate and tropical forest stands, and between terrestrial and aquatic resources. These differences enable exploration of the relative importance of the food resources, as well as movement and dispersal of insects across habitats. The ¹³C‐enrichment (approximately 3‰) caused by saprotrophic fungi can allow the estimation of the importance of fungi in insect diets. Stable N isotopes have revealed food resource partitioning across diverse insect species above and belowground. Detritivorous insects often show a large trophic enrichment in ¹³C (up to 3‰) and ¹⁵N (up to 10‰) relative to the food substrates, soil organic matter. These values are greater than those commonly used for estimation of trophic level. This enrichment likely reflects the prevalence of soil microbial processes, such as fungal development and humification, influencing the isotopic signatures of diets in detritivores. Isotope analysis can become an essential tool in the exploration of insect trophic ecology in terms of biogeochemical C and N cycles, including trophic interactions, plant physiological and soil microbial processes.     

Towards a mechanistic understanding of temperature and enrichment effects on species interaction strength,omnivory and food‐web structure

Revealing the links between species functional traits, interaction strength and food‐web structure is of paramount importance for understanding and predicting the relationships between food‐web diversity and stability in a rapidly changing world. However, little is known about the interactive effects of environmental perturbations on individual species, trophic interactions and ecosystem functioning. Here, we combined modelling and laboratory experiments to investigate the effects of warming and enrichment on a terrestrial tritrophic system. We found that the food‐web structure is highly variable and switches between exploitative competition and omnivory depending on the effects of temperature and enrichment on foraging behaviour and species interaction strength. Our model contributes to identifying the mechanisms that explain how environmental effects cascade through the food web and influence its topology. We conclude that considering environmental factors and flexible food‐web structure is crucial to improve our ability to predict the impacts of global changes on ecosystem diversity and stability.     

Evolution of dispersal polymorphism and local adaptation of dispersal distance in spatially structured landscapes

Many organisms show polymorphism in dispersal distance strategies. This variation is particularly ecological relevant if it encompasses a functional separation of short‐ (SDD) and long‐distance dispersal (LDD). It remains, however, an open question whether both parts of the dispersal kernel are similarly affected by landscape related selection pressures. We implemented an individual‐based model to analyze the evolution of dispersal traits in fractal landscapes that vary in the proportion of habitat and its spatial configuration. Individuals are parthenogenetic with dispersal distance determined by two alleles on each individual's genome: one allele coding for the probability of global dispersal and one allele coding for the variance σ of a Gaussian local dispersal with mean value zero. Simulations show that mean distances of local dispersal and the probability of global dispersal, increase with increasing habitat availability, but that changes in the habitat's spatial autocorrelation impose opposing selective pressure: local dispersal distances decrease and global dispersal probabilities increase with decreasing spatial autocorrelation of the available habitat. Local adaptation of local dispersal distance emerges in landscapes with less than 70% of clumped habitat. These results demonstrate that long and short distance dispersal evolve separately according to different properties of the landscape. The landscape structure may consequently largely affect the evolution of dispersal distance strategies and the level of dispersal polymorphism.     

Confronting the Paradox of Enrichment to the Metacommunity Perspective

Resource enrichment can potentially destabilize predator-prey dynamics. This phenomenon historically referred as the "paradox of enrichment" has mostly been explored in spatially homogenous environments. However, many predator-prey communities exchange organisms within spatially heterogeneous networks called metacommunities. This heterogeneity can result from uneven distribution of resources among communities and thus can lead to the spreading of local enrichment within metacommunities. Here, we adapted the original Rosenzweig-MacArthur predator-prey model, built to study the paradox of enrichment, to investigate the effect of regional enrichment and of its spatial distribution on predator-prey dynamics in metacommunities. We found that the potential for destabilization was depending on the connectivity among communities and the spatial distribution of enrichment. In one hand, we found that at low dispersal regional enrichment led to the destabilization of predator-prey dynamics. This destabilizing effect was more pronounced when the enrichment was uneven among communities. In the other hand, we found that high dispersal could stabilize the predator-prey dynamics when the enrichment was spatially heterogeneous. Our results illustrate that the destabilizing effect of enrichment can be dampened when the spatial scale of resource enrichment is lower than that of organismss movements (heterogeneous enrichment). From a conservation perspective, our results illustrate that spatial heterogeneity could decrease the regional extinction risk of species involved in specialized trophic interactions. From the perspective of biological control, our results show that the heterogeneous distribution of pest resource could favor or dampen outbreaks of pests and of their natural enemies, depending on the spatial scale of heterogeneity.     

Epidemiological landscape models reproduce cyclic insect outbreaks

Forest insect outbreaks can have large impacts on ecosystems and understanding the underlying ecological processes is critical for their management. Current process-based modeling approaches of insect outbreaks are often based on population processes operating at small spatial scales (i.e. within individual forest stands). As such, they are difficult to parameterize and offer limited applicability when modeling and predicting outbreaks at the landscape level where management actions take place. In this paper, we propose a new process-based landscape model of forest insect outbreaks that is based on stand defoliation, the Forest-Infected-Recovering-Forest (FIRF) model. We explore both spatially-implicit (mean field equations with global dispersal) and spatially-explicit (cellular automata with limited dispersal between neighboring stands) versions of this model to assess the role of dispersal in the landscape dynamics of outbreaks. We show that density-dependent dispersal is necessary to generate cyclic outbreaks in the spatially-implicit version of the model. The spatially-explicit FIRF model with local and stochastic dispersal displays cyclic outbreaks at the landscape scale and patchy outbreaks in space, even without density-dependence. Our simple, process-based FIRF model reproduces large scale outbreaks and can provide an innovative approach to model and manage forest pests at the landscape scale.     

Geographical ecology of the palms (Arecaceae): determinants of diversity and distributions across spatial scales

Background

The palm family occurs in all tropical and sub-tropical regions of the world. Palms are of high ecological and economical importance, and display complex spatial patterns of species distributions and diversity.

Scope

This review summarizes empirical evidence for factors that determine palm species distributions, community composition and species richness such as the abiotic environment (climate, soil chemistry, hydrology and topography), the biotic environment (vegetation structure and species interactions) and dispersal. The importance of contemporary vs. historical impacts of these factors and the scale at which they function is discussed. Finally a hierarchical scale framework is developed to guide predictor selection for future studies.

Conclusions

Determinants of palm distributions, composition and richness vary with spatial scale. For species distributions, climate appears to be important at landscape and broader scales, soil, topography and vegetation at landscape and local scales, hydrology at local scales, and dispersal at all scales. For community composition, soil appears important at regional and finer scales, hydrology, topography and vegetation at landscape and local scales, and dispersal again at all scales. For species richness, climate and dispersal appear to be important at continental to global scales, soil at landscape and broader scales, and topography at landscape and finer scales. Some scale–predictor combinations have not been studied or deserve further attention, e.g. climate on regional to finer scales, and hydrology and topography on landscape and broader scales. The importance of biotic interactions – apart from general vegetation structure effects – for the geographic ecology of palms is generally underexplored. Future studies should target scale–predictor combinations and geographic domains not studied yet. To avoid biased inference, one should ideally include at least all predictors previously found important at the spatial scale of investigation.     

Prey dispersal and predator persistence

To understand how patchiness influences population dynamics of a tri-trophic interaction, a tractable model is formulated in terms of differential equations. Motivated by the structure of systems such as plants, phytophagous mites and predatory mites, the model takes dispersal into account at the middle trophic level. The effect of dispersal for the middle level in a tri-trophic system could be either stabilising or destabilising since the middle level acts both as prey and as predator. First a simple model with logistic growth for the lowest level is formulated. A model with logistic growth for the lowest level and instantaneous dispersal has a globally stable three-species equilibrium, if this equilibrium exists. Addition of a middle level dispersal phase of non-negligible duration influences equilibrium stability. In the absence of the top trophic level a limit cycle can occur, caused by the delay that exists in the reaction of the middle level to the changes in the lowest level. With low predator efficiency, it is also possible to have an unstable three-species equilibrium. So addition of a middle level dispersal phase of non-negligible duration can work destabilising. Next the persistence of the third trophic level is studied. Even when the three-species equilibrium exists, the third trophic level need not be persistent. A two-species limit cycle can keep its stability when a three-species equilibrium exists; the system is then bistable. It is argued that such a bistability can offer an alternative explanation for pesticide-induced outbreaks of spider mites and failure of predator introduction.     

Persistence and diversity of directional landscape connectivity improves biomass pulsing in simulations of expanding and contracting wetlands

In flood-pulsed ecosystems, hydrology and landscape structure mediate transfers of energy up the food chain by expanding and contracting in area, enabling spatial expansion and growth of fish populations during rising water levels, and subsequent concentration during the drying phase. Connectivity of flooded areas is dynamic as waters rise and fall, and is largely determined by landscape geomorphology and anisotropy. We developed a methodology for simulating fish dispersal and concentration on spatially-explicit, dynamic floodplain wetlands with pulsed food web dynamics, to evaluate how changes in connectivity through time contribute to the concentration of fish biomass that is essential for higher trophic levels. The model also tracks a connectivity index (DCI) over different compass directions to see if fish biomass dynamics can be related in a simple way to topographic pattern. We demonstrate the model for a seasonally flood-pulsed, oligotrophic system, the Everglades, where flow regimes have been greatly altered. Three dispersing populations of functional fish groups were simulated with empirically-based dispersal rules on two landscapes, and two twelve-year time series of managed water levels for those areas were applied. The topographies of the simulations represented intact and degraded ridge-and-slough landscapes (RSL). Simulation results showed large pulses of biomass concentration forming during the onset of the drying phase, when water levels were falling and fish began to converge into the sloughs. As water levels fell below the ridges, DCI declined over different directions, closing down dispersal lanes, and fish density spiked. Persistence of intermediate levels of connectivity on the intact RSL enabled persistent concentration events throughout the drying phase. The intact landscape also buffered effects of wet season population growth. Water level reversals on both landscapes negatively affected fish densities by depleting fish populations without allowing enough time for them to regenerate. Testable, spatiotemporal predictions of the timing, location, duration, and magnitude of fish concentration pulses were produced by the model, and can be applied to restoration planning.