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。     

Spreading speeds and traveling waves in competitive recursion systems

This paper is concerned with the spreading speeds and traveling wave solutions of discrete time recursion systems, which describe the spatial propagation mode of two competitive invaders. We first establish the existence of traveling wave solutions when the wave speed is larger than a given threshold. Furthermore, we prove that the threshold is the spreading speed of one species while the spreading speed of the other species is distinctly slower compared to the case when the interspecific competition disappears. Our results also show that the interspecific competition does affect the spread of both species so that the eventual population densities at the coexistence domain are lower than the case when the competition vanishes.     

A metric of influential spreading during contagion dynamics through the air transportation network

The spread of infectious diseases at the global scale is mediated by long-range human travel. Our ability to predict the impact of an outbreak on human health requires understanding the spatiotemporal signature of early-time spreading from a specific location. Here, we show that network topology, geography, traffic structure and individual mobility patterns are all essential for accurate predictions of disease spreading. Specifically, we study contagion dynamics through the air transportation network by means of a stochastic agent-tracking model that accounts for the spatial distribution of airports, detailed air traffic and the correlated nature of mobility patterns and waiting-time distributions of individual agents. From the simulation results and the empirical air-travel data, we formulate a metric of influential spreading--the geographic spreading centrality--which accounts for spatial organization and the hierarchical structure of the network traffic, and provides an accurate measure of the early-time spreading power of individual nodes.     

The Speed of Range Shifts in Fragmented Landscapes

Species may be driven extinct by climate change, unless their populations are able to shift fast enough to track regions of suitable climate. Shifting will be faster as the proportion of suitable habitat in the landscape increases. However, it is not known how the spatial arrangement of habitat will affect the speed of range advance, especially when habitat is scarce, as is the case for many specialist species. We develop methods for calculating the speed of advance that are appropriate for highly fragmented, stochastic systems. We reveal that spatial aggregation of habitat tends to reduce the speed of advance throughout a wide range of species parameters: different dispersal distances and dispersal kernel shapes, and high and low extinction probabilities. In contrast, aggregation increases the steady-state proportion of habitat that is occupied (without climate change). Nonetheless, we find that it is possible to achieve both rapid advance and relatively high patch occupancy when the habitat has a “channeled” pattern, resembling corridors or chains of stepping stones. We adapt techniques from electrical circuit theory to predict the rate of advance efficiently for complex, realistic landscape patterns, whereas the rate cannot be predicted by any simple statistic of aggregation or fragmentation. Conservationists are already advocating corridors and stepping stones as important conservation tools under climate change, but they are vaguely defined and have so far lacked a convincing basis in fundamental population biology. Our work shows how to discriminate properties of a landscape''s spatial pattern that affect the speed of colonization (including, but not limited to, patterns like corridors and chains of stepping stones), and properties that affect a species'' probability of persistence once established. We can therefore point the way to better land use planning approaches, which will provide functional habitat linkages and also maintain local population viability.     

Existence of Traveling Waves for the Generalized F–KPP Equation

Variation in genotypes may be responsible for differences in dispersal rates, directional biases, and growth rates of individuals. These traits may favor certain genotypes and enhance their spatiotemporal spreading into areas occupied by the less advantageous genotypes. We study how these factors influence the speed of spreading in the case of two competing genotypes under the assumption that spatial variation of the total population is small compared to the spatial variation of the frequencies of the genotypes in the population. In that case, the dynamics of the frequency of one of the genotypes is approximately described by a generalized Fisher–Kolmogorov–Petrovskii–Piskunov (F–KPP) equation. This generalized F–KPP equation with (nonlinear) frequency-dependent diffusion and advection terms admits traveling wave solutions that characterize the invasion of the dominant genotype. Our existence results generalize the classical theory for traveling waves for the F–KPP with constant coefficients. Moreover, in the particular case of the quadratic (monostable) nonlinear growth–decay rate in the generalized F–KPP we study in detail the influence of the variance in diffusion and mean displacement rates of the two genotypes on the minimal wave propagation speed.     

Spreading speeds of spatially periodic integro-difference models for populations with nonmonotone recruitment functions

An idea used by Thieme (J. Math. Biol. 8, 173-187, 1979) is extended to show that a class of integro-difference models for a periodically varying habitat has a spreading speed and a formula for it, even when the recruitment function R(u, x) is not nondecreasing in u, so that overcompensation occurs. Numerical simulations illustrate the behavior of solutions of the recursion whose initial values vanish outside a bounded set.     

Keeping pace with climate change: what can we learn from the spread of Lessepsian migrants?

Species need to move to keep pace with changing climates, but we do not know if species can move at the required speed. Spread rates of native species may underestimate how fast species can move, we therefore assessed how fast Lessepsian species (marine non‐native species that invaded the Mediterranean from the Red Sea through the Suez Canal) can spread to give a ‘best‐case’ assessment of the effects of climate change on marine biodiversity. We show that about 20% of Lessepsian species could not spread fast enough to keep pace with climate change in about 20% of the global seas and this suggests that climate change may lead to biodiversity loss. The velocity of climate change on the seabed is much lower than at the sea surface, and as a result of this the proportion of species that keep pace with climate change at the seabed was much larger than at the sea surface. This suggests that locations at depth could act as refuges for slow dispersing species. Our analysis compared different estimates of the spreading abilities of marine species and showed that the estimate of spread rates strongly affects the predicted effect of climate change on biodiversity. Providing more accurate estimates of the spreading ability of marine species should therefore have priority if we want to predict the effect of climate change on marine biodiversity. This study is a first approximation of the potential scale and distribution of global marine biodiversity loss and can provide benchmark estimates of the spread rates that species could achieve in colonizing suitable habitat. Assisted colonization may be required to maintain biodiversity in the most strongly affected areas.     

Wind effects on habitat distributions of wind-dispersed invasive plants across different biomes on a global scale: Assessment using six species

A number of widespread invasive plants are wind-dispersed, and wind may facilitate their dispersal and migration over a large distance. While wind is an important factor for seed dispersal and pollination, few studies have examined its potential to affect the habitat distribution of invasive plants over large spatial scales. We selected six of the world's worst invasive plants with wind-driving seed dispersal and pollination, and used wind speed as an indicator of wind. Environmental niche modelling was used to quantify the effects of wind on the habitat distribution of these invasive plants on a global scale and across 14 biomes. Wind had a negative effect on the habitat distribution of invasive plants in tropical and subtropical moist biomes, and a positive effect in Temperate Conifer Forests, Boreal Forests/Taiga, Temperate Grasslands, Savannas and Shrublands, and Montane Grasslands and Shrublands. We concluded that wind affected the habitat distribution of wind-dispersed invasive plants over a large scale, and this effect varied across different biomes. Thus, wind speed and biomes should be used as global monitoring indicators of invasion by wind-dispersed plants and wind speed variables should be included in the projection of habitat distributions of such invasive species when using ENM.     

On interacting populations that disperse to avoid crowding: The effect of a sedentary colony

An analysis is given of a model for two interacting species, one mobile and the other sedentary, in which the mobile one disperses to avoid crowding. The spatial distribution of the mobile species over the habitat, as it evolves with time, is studied. In particular it is shown that a colony of the sedentary species can form an effective barrier against the spreading mobile species, and prevent it from entering certain parts of the habitat.     

Mathematical analysis of the optimal habitat configurations for species persistence

We study a reaction-diffusion model in a binary environment made of habitat and non-habitat regions. Environmental heterogeneity is expressed through the species intrinsic growth rate coefficient. It was known that, for a fixed habitat abundance, species survival depends on habitat arrangements. Our goal is to describe the spatial configurations of habitat that maximise the chances of survival. Through numerical computations, we find that they are of two main types - ball-shaped or stripe-shaped. We formally prove that these optimal shapes depend on the habitat abundance and on the amplitude of the growth rate coefficient. We deduce from these observations that the optimal shape of the habitat realises a compromise between reducing the detrimental habitat edge effects and taking advantage of the domain boundary effects. In the case of an infinite-periodic environment, we prove that the optimal habitat shapes can be deduced from those in the case of a bounded domain.     

Managing mature forest features: The production,accuracy and ecological relevance of a landscape‐scale map

Mature trees and forests contain structural features such as tree hollows, large coarse woody debris and large spreading crowns that provide critical habitat for a wide range of species. These features can take hundreds of years to develop and require careful management to ensure their continued availability. Managing these features requires spatial mapping layers to facilitate landscape‐scale management. This paper outlines how a map of mature forest habitat was developed for Tasmania, Australia. The map was produced using spatial data on vegetation type, mature crown density and senescence, a global layer of forest loss data derived from satellite imagery, a database on timber harvest plans and a spatial layer on the extent of fire. The relationship between mapped mature habitat availability (high, medium, low or negligible) and tree hollow availability in wet forest areas was explored, complementing a similar published study in dry forests. The number of large trees likely to have hollows significantly increased with mapped mature habitat availability, although there was considerable variation and overlap between map categories. Data from a fauna locality database and two radio‐tracking studies showed that three of the vertebrate hollow‐using species examined (Swift Parrot, Common Brushtail Possum and the Tasmanian Long‐eared Bat) and nest records of a species reliant on large tree crowns (the Wedge‐tailed Eagle) were all more likely to occur in areas of higher mapped mature habitat availability. It is concluded that this map reflects the relative availability of tree hollows, is ecologically meaningful and will be useful when managing mature forest habitat at large spatial scales, but the variable accuracy of the map at fine scales needs to be taken into account.     

Present and future projections of habitat suitability of the Asian tiger mosquito,a vector of viral pathogens,from global climate simulation

Climate change can influence the transmission of vector-borne diseases (VBDs) through altering the habitat suitability of insect vectors. Here we present global climate model simulations and evaluate the associated uncertainties in view of the main meteorological factors that may affect the distribution of the Asian tiger mosquito (Aedes albopictus), which can transmit pathogens that cause chikungunya, dengue fever, yellow fever and various encephalitides. Using a general circulation model at 50 km horizontal resolution to simulate mosquito survival variables including temperature, precipitation and relative humidity, we present both global and regional projections of the habitat suitability up to the middle of the twenty-first century. The model resolution of 50 km allows evaluation against previous projections for Europe and provides a basis for comparative analyses with other regions. Model uncertainties and performance are addressed in light of the recent CMIP5 ensemble climate model simulations for the RCP8.5 concentration pathway and using meteorological re-analysis data (ERA-Interim/ECMWF) for the recent past. Uncertainty ranges associated with the thresholds of meteorological variables that may affect the distribution of Ae. albopictus are diagnosed using fuzzy-logic methodology, notably to assess the influence of selected meteorological criteria and combinations of criteria that influence mosquito habitat suitability. From the climate projections for 2050, and adopting a habitat suitability index larger than 70%, we estimate that approximately 2.4 billion individuals in a land area of nearly 20 million km² will potentially be exposed to Ae. albopictus. The synthesis of fuzzy-logic based on mosquito biology and climate change analysis provides new insights into the regional and global spreading of VBDs to support disease control and policy making.     

Fine‐scale foraging habitat selection by two diving central place foragers in the Northeast Atlantic

Habitat selection and spatial usage are important components of animal behavior influencing fitness and population dynamic. Understanding the animal–habitat relationship is crucial in ecology, particularly in developing strategies for wildlife management and conservation. As this relationship is governed by environmental features and intra‐ and interspecific interactions, habitat selection of a population may vary locally between its core and edges. This is particularly true for central place foragers such as gray and harbor seals, where, in the Northeast Atlantic, the availability of habitat and prey around colonies vary at local scale. Here, we study how foraging habitat selection may vary locally under the influence of physical habitat features. Using GPS/GSM tags deployed at different gray and harbor seals’ colonies, we investigated spatial patterns and foraging habitat selection by comparing trip characteristics and home‐range similarities and fitting GAMMs to seal foraging locations and environmental data. To highlight the importance of modeling habitat selection at local scale, we fitted individual models to colonies as well as a global model. The global model suffered from issues of homogenization, while colony models showed that foraging habitat selection differed markedly between regions for both species. Despite being capable of undertaking far‐ranging trips, both gray and harbor seals selected their foraging habitat depending on local availability, mainly based on distance from the last haul‐out and bathymetry. Distance from shore and tidal current also influenced habitat preferences. Results suggest that local conditions have a strong influence on population spatial ecology, highlighting the relevance of processes occurring at fine geographical scale consistent with management within regional units.     

The Modeling of Global Epidemics: Stochastic Dynamics and Predictability

The global spread of emergent diseases is inevitably entangled with the structure of the population flows among different geographical regions. The airline transportation network in particular shrinks the geographical space by reducing travel time between the world's most populated areas and defines the main channels along which emergent diseases will spread. In this paper, we investigate the role of the large-scale properties of the airline transportation network in determining the global propagation pattern of emerging diseases. We put forward a stochastic computational framework for the modeling of the global spreading of infectious diseases that takes advantage of the complete International Air Transport Association 2002 database complemented with census population data. The model is analyzed by using for the first time an information theory approach that allows the quantitative characterization of the heterogeneity level and the predictability of the spreading pattern in presence of stochastic fluctuations. In particular we are able to assess the reliability of numerical forecast with respect to the intrinsic stochastic nature of the disease transmission and travel flows. The epidemic pattern predictability is quantitatively determined and traced back to the occurrence of epidemic pathways defining a backbone of dominant connections for the disease spreading. The presented results provide a general computational framework for the analysis of containment policies and risk forecast of global epidemic outbreaks. On leave from CEA-Centre d'Etudes de Bruyères-Le-Chatel, France.     

A general model of local competition for space

Local competition for space across a wide array of taxa typically involves three mechanisms that we denote here as expansion (spreading into unoccupied habitat), lottery (replacing dead competitors), and overgrowth (encroaching on competitors along zones of contact). By formulating and analysing a simple, general model incorporating these features, we identify ecological conditions and life‐history features that lead to stable coexistence or competitive exclusion (with or without initial‐condition dependence) and gain insight by linking these to case studies in the literature. We demonstrate the importance of contact inhibition, a little‐studied feature of overgrowth, and we show how life‐history tradeoffs may influence and be influenced by local competition for space. The general model we present can help indicate whether local interactions are sufficient to explain patterns of coexistence or exclusion and can serve as the foundation for more specific, realistic models of spatial competition.     

Contrasting spatial and temporal global change impacts on butterfly species richness during the 20th century

Regional patterns of species richness are often explained by models using temperature or measures habitat suitability. Generally, species richness is positively associated with temperature, and negatively associated with habitat degradation. While these models have been well tested across spatial scales, they have rarely been tested on a temporal scale – in part due to the difficulty in ascertaining accurate historical data at an appropriate resolution. In this study, we compared the results of temporal and spatial models, each incorporating two predictors of species richness: temperature, and human population density (as a surrogate of human-related habitat impacts). We found that the change in species richness from the early to late part of the 20th century was positively correlated with temperature change, and negatively correlated with human population density change. When we compared these results to two spatial models using contemporary and historic data, the spatial effects of temperature on butterfly richness were similar to its temporal effects, while the effect of human population density through time is the opposite of its spatial effect. More generally, the assumption that spatial patterns are equivalent to temporal ones when applying macroecological data to global change is clearly unreliable.     

Comparing the consequences of natural selection,adaptive phenotypic plasticity,and matching habitat choice for phenotype–environment matching,population genetic structure,and reproductive isolation in meta‐populations

Organisms commonly experience significant spatiotemporal variation in their environments. In response to such heterogeneity, different mechanisms may act that enhance ecological performance locally. However, depending on the nature of the mechanism involved, the consequences for populations may differ greatly. Building on a previous model that investigated the conditions under which different adaptive mechanisms (co)evolve, this study compares the ecological and evolutionary population consequences of three very different responses to environmental heterogeneity: matching habitat choice (directed gene flow), adaptive plasticity (associated with random gene flow), and divergent natural selection. Using individual‐based simulations, we show that matching habitat choice can have a greater adaptive potential than plasticity or natural selection: it allows for local adaptation while protecting genetic polymorphism despite global mating or strong environmental changes. Our simulations further reveal that increasing environmental fluctuations and unpredictability generally favor the emergence of specialist genotypes but that matching habitat choice is better at preventing local maladaptation by individuals. This confirms that matching habitat choice can speed up the genetic divergence among populations, cause indirect assortative mating via spatial clustering, and hence even facilitate sympatric speciation. This study highlights the potential importance of directed dispersal in local adaptation and speciation, stresses the difficulty of deriving its operation from nonexperimental observational data alone, and helps define a set of ecological conditions which should favor its emergence and subsequent detection in nature.     

Dispersal: risk spreading versus local adaptation

I investigate how risk spreading in stochastic environments and adaptation to permanent properties of local habitats interplay in the simultaneous evolution of dispersal and habitat specialization. In a simple two-patch model, I find many types of locally evolutionarily stable attractors of dispersal and of a trait involved in habitat specialization, including a single habitat specialist and a coalition of two specialists with low dispersal, a generalist with high dispersal, and several types of dispersal polymorphisms. In general, only one attractor is a global evolutionarily stable strategy (ESS). In addition to the ESS analysis, I also present some examples of the dynamics of evolution that exhibit adaptive diversification by evolutionary branching.     

Environmental and topographic drivers of amphibian phylogenetic diversity and endemism in the Iberian Peninsula

Understanding the ecological and evolutionary processes driving biodiversity patterns and allowing their persistence is of utmost importance. Many hypotheses have been proposed to explain spatial diversity patterns, including water-energy availability, habitat heterogeneity, and historical climatic refugia. The main goal of this study is to identify if general spatial drivers of species diversity patterns of phylogenetic diversity (PD) and phylogenetic endemism (PE) at the global scale are also predictive of PD and PE at regional scales, using Iberian amphibians as a case study. Our main hypothesis assumes that topography along with contemporary and historical climate are drivers of phylogenetic diversity and endemism, but that the strength of these predictors may be weaker at the regional scale than it tends to be at the global scale. We mapped spatial patterns of Iberian amphibians' phylogenetic diversity and endemism, using previously published phylogenetic and distribution data. Furthermore, we compiled spatial data on topographic and climatic variables related to the water-energy availability, topography, and historical climatic instability hypotheses. To test our hypotheses, we used Spatial Autoregressive Models and selected the best model to explain diversity patterns based on Akaike Information Criterion. Our results show that, out of the variables tested in our study, water-energy availability and historical climate instability are the most important drivers of amphibian diversity in Iberia. However, as predicted, the strength of these predictors in our case study is weaker than it tends to be at global scales. Thus, additional drivers should also be investigated and we suggest caution when interpreting these predictors as surrogates for different components of diversity.     

Selection and application of spatial indicators for nature conservation at different institutional levels

As European integration increasingly affects pan-European nature conservation, indicators for the assessment of habitats are urgently needed to support ecosystem integrity monitoring as well as the target of halting biodiversity loss by 2010. The Natura 2000 network of protected sites with a strong focus on the protection of habitat types and strict monitoring obligations is now legally binding for all Member States. From a set of indicators that have been proposed for habitat monitoring by the SPIN project (Spatial Indicators for European Nature Conservation) we describe measures of landscape structure and soil function and their potential for the monitoring and management of protected areas and the surrounding landscape. In a case study from Austria, we show that structure-related indicators hold potential for the documentation of local-scale changes on a degraded raised bog Natura 2000 site. In a regional scale case study in northern Germany, we show how landscape metrics relate agricultural statistics, e.g. farm size and livestock density to landscape structure. In a third case study from Slovenia, we show how coarse-scale soil data can be disaggregated to finer scale by integrating topographic information and additional parameters for modelling, and production of soil-related habitat suitability maps. From these case studies we provide an overview of some of the critical issues affecting the selection and application of spatial indicators for nature conservation monitoring tasks. End users of spatial indicators work at different scales and in different biogeographical regions. The indicator selection and application demonstrated in our three case studies reveals the capability to contribute to a more quantitative evidence base for monitoring and management of biodiversity in Europe.