Demographic consequences of population subdivision on the long-furred woolly mouse opossum (Micoureus paraguayanus) from the Atlantic Forest

Habitat destruction and fragmentation severely affected the Atlantic Forest. Formerly contiguous populations may become subdivided into a larger number of smaller populations, threatening their long-term persistence. The computer package VORTEX was used to simulate the consequences of habitat fragmentation and population subdivision on Micoureus paraguayanus, an endemic arboreal marsupial of the Atlantic Forest. Scenarios simulated hypothetical populations of 100 and 2000 animals being partitioned into 1–10 populations, linked by varying rates of inter-patch dispersal, and also evaluated male-biased dispersal. Results demonstrated that a single population was more stable than an ensemble of populations of equal size, irrespective of dispersal rate. Small populations (10–20 individuals) exhibited high instability due to demographic stochasticity, and were characterized by high rates of extinction, smaller values for metapopulation growth and larger fluctuations in population size and growth rate. Dispersal effects on metapopulation persistence were related to the size of the populations and to the sexes that were capable of dispersing. Male-biased dispersal had no noticeable effects on metapopulation extinction dynamics, whereas scenarios modelling dispersal by both sexes positively affected metapopulation dynamics through higher growth rates, smaller fluctuations in growth rate, larger final metapopulation sizes and lower probabilities of extinction. The present study highlights the complex relationships between metapopulation size, population subdivision, habitat fragmentation, rate of inter-patch dispersal and sex-biased dispersal and indicates the importance of gaining a better understanding of dispersal and its interactions with correlations between disturbance events.     

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.     

Local and Regional Determinants of Colonisation-Extinction Dynamics of a Riparian Mainland-Island Root Vole Metapopulation

The role of local habitat geometry (habitat area and isolation) in predicting species distribution has become an increasingly more important issue, because habitat loss and fragmentation cause species range contraction and extinction. However, it has also become clear that other factors, in particular regional factors (environmental stochasticity and regional population dynamics), should be taken into account when predicting colonisation and extinction. In a live trapping study of a mainland-island metapopulation of the root vole (Microtus oeconomus) we found extensive occupancy dynamics across 15 riparian islands, but yet an overall balance between colonisation and extinction over 4 years. The 54 live trapping surveys conducted over 13 seasons revealed imperfect detection and proxies of population density had to be included in robust design, multi-season occupancy models to achieve unbiased rate estimates. Island colonisation probability was parsimoniously predicted by the multi-annual density fluctuations of the regional mainland population and local island habitat quality, while extinction probability was predicted by island population density and the level of the recent flooding events (the latter being the main regionalized disturbance regime in the study system). Island size and isolation had no additional predictive power and thus such local geometric habitat characteristics may be overrated as predictors of vole habitat occupancy relative to measures of local habitat quality. Our results suggest also that dynamic features of the larger region and/or the metapopulation as a whole, owing to spatially correlated environmental stochasticity and/or biotic interactions, may rule the colonisation – extinction dynamics of boreal vole metapopulations. Due to high capacities for dispersal and habitat tracking voles originating from large source populations can rapidly colonise remote and small high quality habitat patches and re-establish populations that have gone extinct due to demographic (small population size) and environmental stochasticity (e.g. extreme climate events).     

Ecological dynamics of extinct species in empty habitat networks. 1. The role of habitat pattern and quantity, stochasticity and dispersal

We examined a remnant host plant ( Primula veris L.) habitat network that was last inhabited by the rare butterfly Hamearis lucina L. in north Wales in 1943, to assess the relative contribution of several spatial parameters to its regional extinction. We first examined relationships between P. veris characteristics and H. lucina eggs in surviving H. lucina populations, and used these to predict the suitability and potential carrying capacity of the habitat network in north Wales. This resulted in an estimate of roughly 4500 eggs (ca 227 adults). We developed a discrete space, discrete time metapopulation model to evaluate the relative contribution of dispersal distance, habitat and environmental stochasticity as possible causes of extinction. We simulated the potential persistence of the butterfly in the current network as well as in three artificial (historical and present) habitat networks that differed in the quantity (current and X3) and fragmentation of the habitat (current and aggregated). We identified that reduced habitat quantity and increased isolation would have increased the probability of regional extinction, in conjunction with environmental stochasticity and H. lucina 's dispersal distance. This general trend did not change in a qualitative manner when we modified the ability of dispersing females to stay in, and find suitable habitats (by changing the size of the grid cells used in the model). Contrary to most metapopulation model predictions, system persistence declined with increasing migration rate, suggesting that the mortality of migrating individuals in fragmented landscapes may pose significant risks to system-wide persistence. Based on model predictions for the present landscape we argue that a major programme of habitat restoration would be required for a re-established metapopulation to persist for >100 years.     

Spatial heterogeneity in the effects of climate and density-dependence on dispersal in a house sparrow metapopulation

Dispersal plays a key role in the response of populations to climate change and habitat fragmentation. Here, we use data from a long-term metapopulation study of a non-migratory bird, the house sparrow (Passer domesticus), to examine the influence of increasing spring temperature and density-dependence on natal dispersal rates and how these relationships depend on spatial variation in habitat quality. The effects of spring temperature and population size on dispersal rate depended on the habitat quality. Dispersal rate increased with temperature and population size on poor-quality islands without farms, where house sparrows were more exposed to temporal fluctuations in weather conditions and food availability. By contrast, dispersal rate was independent of spring temperature and population size on high-quality islands with farms, where house sparrows had access to food and shelter all the year around. This illustrates large spatial heterogeneity within the metapopulation in how population density and environmental fluctuations affect the dispersal process.     

Effect of habitat fragmentation on dispersal in the butterfly Proclossiana eunomia

Comparison of dispersal rates of the bog fritillary butterfly between continuous and fragmented landscapes indicates that between patch dispersal is significantly lower in the fragmented landscape, while population densities are of the same order of magnitude. Analyses of the dynamics of the suitable habitat for the butterfly in the fragmented landscape reveal a severe, non linear increase in spatial isolation of patches over a time period of 30 years (i.e. 30 butterfly generations), but simulations of the butterfly metapopulation dynamics using a structured population model show that the lower dispersal rates in the fragmented landscape are far above the critical threshold leading to metapopulation extinction. These results indicate that changes in individual behaviour leading to the decrease of dispersal rates in the fragmented landscape were rapidly selected for when patch spatial isolation increased. The evidence of such an adaptive answer to habitat fragmentation suggests that dispersal mortality is a key factor for metapopulation persistence in fragmented landscapes. We emphasise that landscape spatial configuration and patch isolation have to be taken into account in the debate about large-scale conservation strategies.     

Temporal autocorrelation can enhance the persistence and abundance of metapopulations comprised of coupled sinks

In spatially heterogeneous landscapes, some habitats may be persistent sources, providing immigrants to sustain populations in unfavorable sink habitats (where extinction is inevitable without immigration). Recent theoretical and empirical studies of source-sink systems demonstrate that temporally variable local growth rates in sinks can substantially increase average abundance of a persisting population, provided that the variation is positively autocorrelated--in effect, temporal variation inflates average abundance. Here we extend these results to a metapopulation in which all habitat patches are sinks. Using numerical studies of a population with discrete generations (buttressed by analytic results), we show that temporal variation and moderate dispersal can jointly permit indefinite persistence of the metapopulation and that positive autocorrelation both lowers the magnitude of variation required for persistence and increases the average abundance of persisting metapopulations. These effects are weakened--but not destroyed--if variation in local growth rates is spatially synchronized and dispersal is localized. We show that the inflationary effect is robust to a number of extensions of the basic model, including demographic stochasticity and density dependence. Because ecological and environmental processes contributing to temporally variable growth rates in natural populations are typically autocorrelated, these observations may have important implications for species persistence.     

Dispersal ability and host-plant characteristics influence spatial population structure of monophagous beetles

Abstract.  1. Dispersal plays an integral role in determining spatial population structure and, consequently, the long-term survival of many species. Theoretical studies indicate that dispersal increases with population density and decreasing habitat stability. In the case of monophagous insect herbivores, the stability of host-plant populations may influence their spatial population structure.
2. The tallgrass prairie in Iowa, U.S.A. is highly fragmented and most prairie insects face a landscape with fewer habitat patches and smaller host-plant populations than 150 years ago, potentially making dispersal between patches difficult. Some herbivores, however, use native plant species with weedy characteristics that have increased in abundance because of disturbances.
3. Mark–recapture data and presence–absence surveys were used to examine dispersal and spatial population structure of two monophagous beetles with host plants that exhibit different population stability and have responded differently to fragmentation of tallgrass prairie.
4. Chrysochus auratus Fabricius exhibits a patchy population structure and has relatively large dispersal distances and frequencies. Its host plant is variable locally in time and space, but is more abundant than 150 years ago. The other species, Anomoea laticlavia Forster, exhibits a metapopulation or non-equilibrium population structure and has relatively small dispersal distances and frequencies. Its host-plant populations are stable in time and space.
5. The results indicate that dispersal ability of monophagous beetles reflects the life-history dynamics of their host plants, but the spatial population structure exhibited today is strongly influenced by how the host plants have responded to the fragmentation process over both time and space.     

Female-biased dispersal, low female recruitment, unpaired males, and the extinction of small and isolated bird populations

Small and isolated populations are usually assumed to be at a high risk of extinction due to environmental or demographic stochasticity, genetic problems, or too little immigration. In birds, natal dispersal is usually female-biased, but the consequences of such a pattern on vulnerability to extinction of isolated populations has not received much attention before. In this paper I derive predictions as to how female-biased natal dispersal may differentially affect the extinction risk of populations and species with contrasting distributions, migratory behaviours, life histories and mating systems. Female-biased dispersal will lead to male-biased sex ratios in small, isolated or fragmented populations, in particular because recent research has shown that females often have a limited ability to search for mates and may therefore effectively be lost from the breeding population if they disperse into areas empty of conspecifics. I reviewed published studies on birds and found that a high proportion of unpaired males is common in isolated populations or populations in small habitat fragments. Dispersal of females may therefore increase the vulnerability to extinction of small or isolated populations, or populations at the periphery of a species' distribution range. I also predict that vulnerability to extinction should be greater for migratory than for resident species and greater for short-lived than for long-lived species because of differences in the time available for females to locate unpaired males. Further, extinction risk may also be greater for birds than for mammals due to differences in which sex disperses and patterns of parental care. Finally, mating system will also affect vulnerability to extinction when natal dispersal leads to biased sex ratios. I review available evidence for these predictions (e.g. songbird declines in North America) and discuss implications for conservation.     

Inbreeding depresses short and long distance dispersal in three congeneric spiders

Dispersal is one of the most important precopulatory inbreeding avoidance mechanisms and subject to landscape related selection pressures. In small populations, inbreeding within and between populations may strongly affect population dynamics if it reduces fitness and gene‐flow. While inbreeding avoidance is generally considered to be a key evolutionary driver of dispersal, potential effects of inbreeding on the dispersal process, are poorly known. Here, I document how inbreeding within a population, so by mating among relatives, affects the survivorship and the dispersal behaviour of three congeneric spider Erigone species (Araneae: Linyphiidae) that differ in habitat preference and regional rarity. The three species were chosen as a model because they allow the assessment of both long and short distance dispersal motivation (respectively ballooning and rappelling) under laboratory conditions. Inbreeding reduced both long and short distance dispersal modes in the three congeneric species. Because survival was depressed after inbreeding, with a tendency of reduced survival loss in the rare and highly stenotopic species, energetic constraints are likely to be the underlying mechanism. Inbreeding consequently depresses silk‐related dispersal in three related spiders. This may induce an inbreeding depression vortex with important consequences for range expansion and metapopulation dynamics of aerially dispersing species from highly fragmented landscapes.     

Host–parasitoid spatial models: the interplay of demographic stochasticity and dynamics

Host–parasitoid metapopulation models have typically been deterministic models formulated with population numbers as a continuous variable. Spatial heterogeneity in local population abundance is a typical (and often essential) feature of these models and means that, even when average population density is high, some patches have small population sizes. In addition, large temporal population fluctuations are characteristic of many of these models, and this also results in periodically small local population sizes. Whenever population abundances are small, demographic stochasticity can become important in several ways. To investigate this problem, we have reformulated a deterministic, host–parasitoid metapopulation as an integer-based model in which encounters between hosts and parasitoids, and the fecundity of individuals are modelled as stochastic processes. This has a number of important consequences: (1) stochastic fluctuations at small population sizes tend to be amplified by the dynamics to cause massive population variability, i.e. the demographic stochasticity has a destabilizing effect; (2) the spatial patterns of local abundance observed in the deterministic counterpart are largely maintained (although the area of ''spatial chaos'' is extended); (3) at small population sizes, dispersal by discrete individuals leads to a smaller fraction of new patches being colonized, so that parasitoids with small dispersal rates have a greater tendency for extinction and higher dispersal rates have a larger competitive advantage; and (4) competing parasitoids that could coexist in the deterministic model due to spatial segregation cannot now coexist for any combination of parameters.     

The effects of habitat fragmentation on the social kin structure and mating system of the agile antechinus, Antechinus agilis

Habitat fragmentation is one of the major contributors to the loss of biodiversity worldwide. However, relatively little is known about its more immediate impacts on within-patch population processes such as social structure and mating systems, whose alteration may play an important role in extinction risk. We investigated the impacts of habitat fragmentation due to the establishment of an exotic softwood plantation on the social kin structure and breeding system of the Australian marsupial carnivore, Antechinus agilis. Restricted dispersal by males in fragmented habitat resulted in elevated relatedness among potential mates in populations in fragments, potentially increasing the risk of inbreeding. Antechinus agilis nests communally in tree hollows; these nests are important points for social contact between males and females in the mating season. In response to elevated relatedness among potential mates in fragmented habitat, A. agilis significantly avoided sharing nests with opposite-sex relatives in large fragment sites (but not in small ones, possibly due to limited nest locations and small population sizes). Because opposite-sex individuals shared nests randomly with respect to relatedness in unfragmented habitat, we interpreted the phenomenon in fragmented habitat as a precursor to inbreeding avoidance via mate choice. Despite evidence that female A. agilis at high inbreeding risk selected relatively unrelated mates, there was no overall increased avoidance of related mates by females in fragmented habitats compared to unfragmented habitats. Simulations indicated that only dispersal, and not nonrandom mating, contributed to inbreeding avoidance in either habitat context. However, habitat fragmentation did influence the mating system in that the degree of multiple paternity was reduced due to the reduction in population sizes and population connectivity. This, in turn, reduced the number of males available to females in the breeding season. This suggests that in addition to the obvious impacts of reduced recruitment, patch recolonization and increased genetic drift, the isolation of populations in habitat patches may cause changes in breeding behaviour that contribute to the negative impacts of habitat fragmentation.     

Long-term demographic surveys reveal a consistent relationship between average occupancy and abundance within local populations of a butterfly metapopulation

Species distribution models are the tool of choice for large-scale population monitoring, environmental association studies and predictions of range shifts under future environmental conditions. Available data and familiarity of the tools rather than the underlying population dynamics often dictate the choice of specific method – especially for the case of presence–absence data. Yet, for predictive purposes, the relationship between occupancy and abundance embodied in the models should reflect the actual population dynamics of the modelled species. To understand the relationship of occupancy and abundance in a heterogeneous landscape at the scale of local populations, we built a spatio-temporal regression model of populations of the Glanville fritillary butterfly Melitaea cinxia in a Baltic Sea archipelago. Our data comprised nineteen years of habitat surveys and snapshot data of land use in the region. We used variance partitioning to quantify relative contributions of land use, habitat quality and metapopulation covariates. The model revealed a consistent and positive, but noisy relationship between average occupancy and mean abundance in local populations. Patterns of abundance were highly variable across years, with large uncorrelated random variation and strong local population stochasticity. In contrast, the spatio-temporal random effect, habitat quality, population connectivity and patch size explained variation in occupancy, vindicating metapopulation theory as the basis for modelling occupancy patterns in fragmented landscapes. Previous abundance was an important predictor in the occupancy model, which points to a spillover of abundance into occupancy dynamics. While occupancy models can successfully model large-scale population structure and average occupancy, extinction probability estimates for local populations derived from occupancy-only models are overconfident, as extinction risk is dependent on actual, not average, abundance.     

Habitat destruction, environmental catastrophes, and metapopulation extinction

The extinction process of fragmented populations, characterized by a small number of conspecifics inhabiting each patch, is heavily affected by natural and human disturbance. To evaluate the risk of extinction we consider a network of identical patches connected by passive or active dispersal and hosting a finite, discrete number of individuals. We discuss three types of disturbance affecting the metapopulation: permanent loss of habitat patches, erosion of existing patches, and random catastrophes that wipe out the entire population of a patch. Starting from an infinite-dimensional Markov model that fully accounts for demographic stochasticity, we reduce it to finite dimension via moment closure with negative-binomial approximation. The compact models obtained in this way account for the dynamics of the fraction of empty patches, the average number of individuals in occupied patches, and the variance of their distribution. After comparing the performance of these compact models with that of the infinite-dimensional model in the case of no disturbances, we then proceed to computing persistence-extinction boundaries as bifurcation lines of the compact models in the space of demographic and disturbance parameters. We consider bifurcations with respect to demographic and environmental parameters and contrast our results with those of previous theories. We find out that environmental catastrophes increase the risk of extinction for both frequent and infrequent dispersers, while the random loss of patches has a much larger influence on frequent dispersers. This influence can be counterbalanced by active dispersal. Local erosion of habitat fragments has a larger influence on infrequent than on frequent dispersers. We finally discuss the important synergistic effects of disturbances acting simultaneously.     

The effects of habitat fragmentation via forestry plantation establishment on spatial genotypic structure in the small marsupial carnivore, Antechinus agilis

Dispersal is an important influence on species' distributions, patch colonization and population persistence in fragmented habitat. We studied the impacts of habitat fragmentation resulting from establishment of an exotic pine plantation on dispersal of the marsupial carnivore, Antechinus agilis. We applied spatial analyses of individual multilocus microsatellite genotypes and mitochondrial haplotypes to study patterns of gene flow in fragmented habitat and natural habitat 'control' areas, and how this is affected by the spatial dispersion of habitat patches, the presence of corridors and a 'mainland' source of migrants. Spatial analysis of molecular variance and partial Mantel tests confirmed the absence of cryptic barriers to gene flow in continuous habitat, which if present would confound the comparison of genetic structures in fragmented vs. unfragmented habitats. Spatial genotypic structure suggested that although dispersal was male-biased in both habitat types, fragmentation restricted dispersal of males more than that of females and the degree of restriction of male dispersal was dependent on the geographical isolation of the patch. The scale of positive genotypic structure in fragmented habitat was restricted to the two closest patches for females and the three closest patches for males. Our results provide evidence for significantly increased gene flow through habitat corridors relative to that across the matrix and for significantly lower gene flow between 'mainland' unfragmented habitat and habitat patches relative to that within either habitat type, suggesting a behavioural barrier to crossing habitat interfaces.     

Allee效应对具有生境恢复的集合种群的影响

Allee效应与种群的灭绝密切相关,其研究对生态保护和管理至关重要。Allee效应对物种续存是潜在的干扰因素,濒危物种更容易受其影响,可能会增加生存于生境破碎化斑块的濒危物种的死亡风险,因此研究Allee效应对种群的动态和续存的影响是必要的。从包含由生物有机体对环境的修复产生的Allee效应的集合种群模型出发,引入由其他机制形成的Allee效应,建立了常微分动力系统模型和基于网格模型的元胞自动机模型。通过理论分析和计算机模拟表明:(1)强Allee效应不利于具有生境恢复的集合种群的续存;(2)生境恢复有利于种群续存;(3)局部扩散影响了集合种群的空间结构、动态行为和稳定性,生境斑块之间的局部作用将会减缓或消除集合种群的Allee效应,有利于集合种群的续存。     

Metapopulation dynamics: Does it help to have more of the same?

With the interest in conservation biology shifting towards processes from patterns, and to populations from communities, the theory of metapopulation dynamics is replacing the equilibrium theory of island biogeography as the population ecology paradigm in conservation biology. The simplest models of metapopulation dynamics make predictions about the effects of habitat fragmentation - size and isolation of habitat patches - on metapopulation persistence. The simple models may be enriched by considerations of the effects of demographic and environmental stochasticity on the size and extinction probability of local populations. Environmental stochasticity affects populations at two levels: it makes local extinctions more probable, and it also decreases metapopulation persistence time by increasing the correlation of extinction events across populations. Some controversy has arisen over the significance of correlated extinctions, and how they may affect the optimal subdivision of metapopulations to maximize their persistence time.     

The fragmented population structure of the Sardinian chalk hill blue butterfly (Lepidoptera, Lycaenidae)

The endemic Sardinian chalk hill blue butterfly, Polyommatus coridon gennargenti, is considered vulnerable to extinction because of its low genetic variation and restricted distribution. The species also has a fragmented distribution, which follows the patchy distribution pattern of its larval host-plant. A preliminary investigation of the population structure of P. coridon gennargenti was carried out on a small network of four local populations by means of capture–recapture methods. Estimated population sizes and movement rates among the four adjacent local populations suggest that this taxon has a metapopulation structure composed of loosely connected small local populations. Natural fragmentation, isolation, and traditional land use contribute to the vulnerability of P. coridon gennargenti to extinction. Low effective population sizes and restricted movement between habitat patches lead to inbreeding and an increased vulnerability to extinction of this island population.     

Relative importance of host plant patch geometry and habitat quality on the patterns of occupancy,extinction and density of the monophagous butterfly <Emphasis Type="Italic">Iolana iolas</Emphasis>

Habitat fragmentation is a major cause of species rarity and decline because it increases local population extinctions and reduces recolonisation rates of remnant patches. Although two major patch characteristics (area and connectivity) have been used to predict distribution patterns in fragmented landscapes, other factors can affect the occurrence of a species as well as the probability of it becoming extinct. In this paper, we study the spatial structure and dynamics of the butterfly Iolana iolas in a 75-patch network of its host plant (Colutea hispanica) to determine the relative importance of patch area, connectivity and habitat quality characteristics on occupancy, extinction and density over the period 2003–2006. Occupancy in 2003, incidence (proportion of years occupied) and probability of extinction were mostly affected by patch area. Smaller patches were less likely to be occupied because they had a higher probability of extinction, partly due to environmental stochasticity. The density of I. iolas was negatively related to patch area in all study years. Only in 2004 was the density of I. iolas positively influenced by fruit production per plant. Our results suggest that for I. iolas, and probably for other specialist butterflies with clearly delimited resource requirements, metapopulation dynamics can be satisfactorily predicted using only geometric variables because most habitat characteristics are subsumed in patch area. However, this hypothesis should be subject to further testing under diverse environmental conditions to evaluate the extent of its generalisation.     

Metapopulation Persistence in Random Fragmented Landscapes

Habitat destruction and land use change are making the world in which natural populations live increasingly fragmented, often leading to local extinctions. Although local populations might undergo extinction, a metapopulation may still be viable as long as patches of suitable habitat are connected by dispersal, so that empty patches can be recolonized. Thus far, metapopulations models have either taken a mean-field approach, or have modeled empirically-based, realistic landscapes. Here we show that an intermediate level of complexity between these two extremes is to consider random landscapes, in which the patches of suitable habitat are randomly arranged in an area (or volume). Using methods borrowed from the mathematics of Random Geometric Graphs and Euclidean Random Matrices, we derive a simple, analytic criterion for the persistence of the metapopulation in random fragmented landscapes. Our results show how the density of patches, the variability in their value, the shape of the dispersal kernel, and the dimensionality of the landscape all contribute to determining the fate of the metapopulation. Using this framework, we derive sufficient conditions for the population to be spatially localized, such that spatially confined clusters of patches act as a source of dispersal for the whole landscape. Finally, we show that a regular arrangement of the patches is always detrimental for persistence, compared to the random arrangement of the patches. Given the strong parallel between metapopulation models and contact processes, our results are also applicable to models of disease spread on spatial networks.