Habitat fragmentation and extinction thresholds on fractal landscapes

Habitat fragmentation is a potentially critical factor in determining population persistence. In this paper, we explore the effect of fragmentation when the fragmentation follows a fractal pattern. The habitat is divided into patches, each of which is suitable or unsuitable. Suitable patches are either occupied or unoccupied, and change state depending on rates of colonization and local extinction. We compare the behaviour of two models: a spatially implicit patch-occupancy (PO) model and a spatially explicit cellular automaton (CA) model. The PO model has two fixed points: extinction, and a stable equilibrium with a fixed proportion of occupied patches. Global extinction results when habitat destruction reduces the proportion of suitable patches below a critical threshold. The PO model successfully recreates the extinction patterns found in other models. We translated the PO model into a stochastic cellular automaton. Fractal arrangements of suitable and unsuitable patches were used to simulate habitat fragmentation. We found that: (i) a population on a fractal landscape can tolerate more habitat destruction than predicted by the patch-occupancy model, and (ii) the extinction threshold decreases as the fractal dimension of the landscape decreases. These effects cannot be seen in spatially implicit models. Landscape struc-ture plays a vital role in mediating the effects of habitat fragmentation on persistence.     

Mixed effects of habitat fragmentation on species richness and community structure in a microarthropod microecosystem

Abstract.  1. Theory is unclear about the optimal degree of isolation of habitat fragments where the aim is to maximise species richness. In a field-based microecosystem of Collembola and predatory and non-predatory mites, moss patches of the same total area were fragmented to varying degrees. The habitat was left for several months to allow the communities to approach a new state of equilibrium.
2. The species richness (in particular of predatory mites) of a given area of habitat was greater when it was part of a large mainland area than part of an island, in agreement with theory.
3. Conversely, species richness and abundance were largely unaffected by fragmentation of a fixed area of island habitat. In this case, it is suggested here that the advantages of several small patches (e.g. reduced impact of environmental stochasticity, wider range of habitats overall) were equally balanced by the advantages of a single large patch (e.g. reduced effect of demographic stochasticity, wider range of habitats within a single patch, reduced edge effect), or that both effects were small.
4. The shapes of rank–abundance curves were similar among the levels of fragmentation of a fixed area of island habitat, implying that fragmentation had little impact on community structure. Conversely, the species composition of non-predatory mites varied weakly, but significantly, by fragmentation.     

Habitat fragmentation resulting in overgrazing by herbivores

Habitat fragmentation sometimes results in outbreaks of herbivorous insect and causes an enormous loss of primary production. It is hypothesized that the driving force behind such herbivore outbreaks is disruption of natural enemy attack that releases herbivores from top-down control. To test this hypothesis I studied how trophic community structure changes along a gradient of habitat fragmentation level using spatially implicit and explicit models of a tri-trophic (plant, herbivore and natural enemy) food chain. While in spatially implicit model number of trophic levels gradually decreases with increasing fragmentation, in spatially explicit model a relatively low level of habitat fragmentation leads to overgrazing by herbivore to result in extinction of the plant population followed by a total system collapse. This provides a theoretical support to the hypothesis that habitat fragmentation can lead to overgrazing by herbivores and suggests a central role of spatial structure in the influence of habitat fragmentation on trophic communities. Further, the spatially explicit model shows (i) that the total system collapse by the overgrazing can occur only if herbivore colonization rate is high; (ii) that with increasing natural enemy colonization rate, the fragmentation level that leads to the system collapse becomes higher, and the frequency of the collapse is lowered.     

Individual-based spatially-explicit model of an herbivore and its resource: the effect of habitat reduction and fragmentation

We present an individual-based, spatially-explicit model of the dynamics of a small mammal and its resource. The life histories of each individual animal are modeled separately. The individuals can have the status of residents or wanderers and belong to behaviorally differing groups of juveniles or adults and males or females. Their territory defending and monogamous behavior is taken into consideration. The resource, green vegetation, grows depending on seasonal climatic characteristics and is diminished due to the herbivore's grazing. Other specifics such as a varying personal energetic level due to feeding and starvation of the individuals, mating preferences, avoidance of competitors, dispersal of juveniles, as a result of site overgrazing, etc., are included in the model. We determined model parameters from real data for the species Microtus ochrogaster (prairie vole). The simulations are done for a case of an enclosed habitat without predators or other species competitors. The goal of the study is to find the relation between size of habitat and population persistence. The experiments with the model show the populations go extinct due to severe overgrazing, but that the length of population persistence depends on the area of the habitat as well as on the presence of fragmentation. Additionally, the total population size of the vole population obtained during the simulations exhibits yearly fluctuations as well as multi-yearly peaks of fluctuations. This dynamics is similar to the one observed in prairie vole field studies.     

Combining endogenous and exogenous spatial variability in analytical population models

Analytically tractable models of dynamics in continuous space rarely incorporate both endogenous and exogenous spatial heterogeneity. We use spatial moment equations in combination with simulation models to analyze the combined effects of endogenous and exogenous variability on population viability in a simple single-population model where landscape heterogeneity and local population density both affect mortality rate. The equations partition the effects of heterogeneity into an effect of local crowding and an effect of habitat association caused by differential mortality. Exogenous heterogeneity in mortality rate increases population viability through habitat association and decreases it through increased crowding; the net effect of exogenous heterogeneity is generally to improve population viability. This result is contrary to some (but not all) conclusions in the literature, which usually focus on the effects of fragmentation rather than the benefits of refuges to short-dispersing individuals.     

Indépendance des formes spectrales de chlorophylle a et des holochromes chlorophylliens

Independence of special forms of chlorophyll a and chlorophyll holochromesZea mays L. seedlings were cultivated for 10 days with submission to 4 s illumination periods interspersed with dark periods varying in length from 30 min to 6 h depending on the lot analyzed. The results show that, for the case in which the dark periods were shorter than 1 h, the relative proportions of different spectroscopic chlorophyll forms (maxima at 662, 670, 677.5, and 684 nm) were constant. For longer durations of darkness between illuminations, the relative proportion of the form Ca₆₇₀ increases, while that of Ca₆₈₄ diminishes with the length of darkness; to a lesser extent, the relative proportion of Ca₆₆₂ increases and a form Ca₆₉₂ disappears. A scheme is proposed to explain the evolution of the relative proportions of the different spectral forms.The different chlorophyll holochromes present in the chloroplasts were also analysed. If the dark period was longer than 1 h, chlorophyll was associated with peptide chains of molecular weights 21 000 and 29 000. If the dark period was shorter than 1 h chlorophyll was associated with four peptide chains of molecular weights 21 000, 25 000, 29 000 and 70 000.The results taken together demonstrate that a given spectral chlorophyll a form cannot be associated with a definite chlorophyll holochrome.     

Disentangling the effects of historic vs. contemporary landscape structure on population genetic divergence

Increasing habitat fragmentation poses an immediate threat to population viability, as gene flow patterns are changed in these altered landscapes. Patterns of genetic divergence can potentially reveal the impact of these shifts in landscape connectivity. However, divergence patterns not only carry the signature of altered contemporary landscapes, but also historical ones. When considered separately, both recent and historical landscape structure appear to significantly affect connectivity among 51 wood frog ( Rana sylvatica ) populations. However, by controlling for correlations among landscape structure from multiple time periods, we show that patterns of genetic divergence reflect recent landscape structure as opposed to landscape structure prior to European settlement of the region (before 1850s). At the same time, within-population genetic diversities remain high and a genetic signature of population bottlenecks is lacking. Together, these results suggest that metapopulation processes – not drift-induced divergence associated with strong demographic bottlenecks following habitat loss – underlie the strikingly rapid consequences of temporally shifting landscape structure on these amphibians. We discuss the implications of these results in the context of understanding the role of population demography in the adaptive variation observed in wood frog populations.     

The measure of order and disorder in the distribution of species in fragmented habitat

Species distribution patterns within naturally fragmented habitat have been found to often exhibit patterns of pronounced nestedness. Highly predictable extinction sequences are implied by these nested species distribution patterns, thus the patterns are important to both the philosophy and practice of conservation biology. A simple thermodynamic measure of the order and disorder apparent in the nested patterns is described. The metric offers (i) a measure of the uncertainty in species extinction order, (ii) a measure of relative populational stabilities, (iii) a means of identifying minimally sustainable population sizes, and (iv) an estimate of the historical coherence of the species assemblage. Four presumptions govern the development of the metric and its theory: (i) the fragmented habitat was once whole and originally populated by a single common source biota, (ii) the islands were initially uniform in their habitat heterogeneity and type mix, and have remained so throughout their post-fragmentation history, (iii) no significant clinal (latitudinal) gradation exists across the archipelago so as to promote species turnover across the archipelago, and (iv) all species of interest are equally isolated on all islands. The violation of these conditions promotes species distributions which are idiosyncratic to the general extinction order expected in fragmentation archipelagos. While some random variation in extinction order is to be expected, idiosyncratic distributional patterns differ from randomness and are readily segregatable from such noise. A method of identifying idiosyncratic species and sites is described.     

The effect of habitat fragmentation on cyclic population dynamics: a reduction to ordinary differential equations

Habitat fragmentation is known to be a key factor affecting population dynamics. In a previous study by Strohm and Tyson (Bull Math Biol 71:1323?C1348, 2009), the effect of habitat fragmentation on cyclic population dynamics was studied using spatially explicit predator?Cprey models with four different sets of reaction terms. The difficulty with spatially explicit models is that often analytical tractability is lost and the mechanisms behind the behaviour of the models are difficult to analyse. In this study, we employ a simplification procedure based on a Fourier series first-term truncation of the spatially explicit models Strohm and Tyson (Bull Math Biol 71:1323?C1348, 2009) to obtain spatially implicit models. These simpler models capture the main features of the spatially explicit models and can be used to explain the dynamics observed by Strohm and Tyson. We find that the spatially implicit models and the spatially explicit models produce similar responses to habitat fragmentation for larger high-quality patch sizes. Additionally, we find that the critical patch size of the spatially implicit models provides an upper bound on the critical patch size of the spatially explicit models. Finally, we derive an approximation of the multi-patch habitat by a single-patch habitat with partial flux boundary conditions which allows for a lower bound on the critical patch size to be calculated.     

Effects of Small Habitat Size and Isolation on the Population Structure of Common Wetland Species

Abstract: Habitat fragmentation may cause plant species to suffer from inbreeding and genetic drift, which affects population viability negatively. Less viable populations may contain an altered population structure, i.e., they have a smaller proportion of seedlings and a larger proportion of vegetative adults, when compared to large extensive populations. We applied a hierarchical-spatial field design, consisting of large habitat islands with surrounding near and distant small habitat islands to distinguish between the two components of habitat fragmentation, i.e., the effect of small habitat size and the effect of isolation. We studied two common habitat-specific species: Carex davalliana (Cyperaceae) and Succisa pratensis (Dipsacaceae). We identified a decrease in the proportion of C. davalliana seedlings and an increase in the proportion of vegetative tillers in response to isolation. In S. pratensis, we identified a decrease in the proportion of seedlings and an increase in the proportion of vegetative rosettes that were attributable to small habitat size. Hence, we confirmed alterations in the population structure of those common species, although both species were affected by a different component of habitat fragmentation. The observed discrepancy between species could be related their life history. We conclude that habitat fragmentation adversely affects not only rare but also common species. Their loss of viability can inevitably lead to alterations in vegetation structure and consequently lead to the further loss of this species-rich and threatened ecosystem type.     

Metapopulation dynamics of the bog fritillary butterfly: modelling the effect of habitat fragmentation

Population viability analysis (PVA) and metapopulation theory are valuable tools to model the dynamics of spatially structured populations. In this article we used a spatially realistic population dynamic model to simulate the trajectory of a Proclossiana eunomia metapopulation in a network of habitat patches located in the Belgian Ardenne. Sensitivity analysis was used to evaluate the relative influence of the different parameters on the model output. We simulated habitat loss by removing a percentage of the original habitat, proportionally in each habitat patch. Additionally, we evaluated isolation and fragmentation effects by removing and dividing habitat patches from the network, respectively. The model predicted a slow decline of the metapopulation size and occupancy. Extinction risks predicted by the model were highly sensitive to environmental stochasticity and carrying capacity. For a determined level of habitat destruction, the expected lifetime of the metapopulation was highly dependent on the spatial configuration of the landscape. Moreover, when the proportion of removed habitat is above 40% of the original habitat, the loss of whole patches invariably leads to the strongest reduction in metapopulation viability.     

Habitat fragmentation: A long and tangled tale

In this essay: I provide a brief history of habitat fragmentation research; I describe why its “non‐questions” (‘Is habitat fragmentation a big problem for wildlife species?” and, “Are the effects of habitat fragmentation generally negative or positive?”) are important to conservation; I outline my role in tackling these questions; I discuss reasons why the culture of habitat fragmentation research is largely incapable of accepting the answers; and I speculate on the future of habitat fragmentation research.     

An island paradigm on the mainland: host population fragmentation impairs the community of avian pathogens

Emergent infectious diseases represent a major threat for biodiversity in fragmented habitat networks, but their dynamics in host metapopulations remain largely unexplored. We studied a large community of pathogens (including 26 haematozoans, bacteria and viruses as determined through polymerase chain reaction assays) in a highly fragmented mainland bird metapopulation. Contrary to recent studies, which have established that the prevalence of pathogens increase with habitat fragmentation owing to crowding and habitat-edge effects, the analysed pathogen parameters were neither dependent on host densities nor related to the spatial structure of the metapopulation. We provide, to our knowledge, the first empirical evidence for a positive effect of host population size on pathogen prevalence, richness and diversity. These new insights into the interplay between habitat fragmentation and pathogens reveal properties of a host-pathogen system resembling island environments, suggesting that severe habitat loss and fragmentation could lower pathogen pressure in small populations.     

Non-linear elastic deformable models for real-time surgery simulation

In this article, we describe the latest developments of the minimally invasive hepatic surgery simulator prototype developed at INRIA. A key problem with such a simulator is the physical modelling of soft tissues. We propose a new deformable model based on non-linear elasticity and the finite element method. This model is valid for large displacements, which means in particular that it is invariant with respect to rotations. This property improves the realism of the deformations and solves the problems related to the shortcomings of linear elasticity, which is only valid for small displacements. We also address the problem of volume variations by adding to our model incompressibility constraints. Finally, we demonstrate the relevance of this approach for the real-time simulation of laparoscopic surgical gestures on the liver.     

广西黑叶猴栖息地景观格局破碎化分析及其对种群的影响

黑叶猴(Trachypithecus francoisi)是仅分布于喀斯特石山生境的珍稀濒危灵长类动物。由于非法捕杀和人类活动干扰,其种群数量正在急剧减少。同时,随着森林砍伐和土地开垦的加速,其栖息地严重破碎化。因此,了解栖息地破碎化对黑叶猴种群的影响对于保护这一珍稀濒危物种具有重要意义。基于遥感影像、土地利用数据以及黑叶猴种群调查数据,通过Fragstats软件开展广西黑叶猴栖息地景观破碎化分析,并通过相关性和多元逐步回归分析,探讨了景观格局对广西黑叶猴种群数量的影响。结果表明：(1)广西黑叶猴栖息地呈现破碎化严峻、斑块形状复杂化、斑块团聚程度较弱且分散化的现象;栖息地以林地景观占据重要优势,但人为景观的干扰十分强烈;在不同地区中,生境破碎化程度、人为干扰强度以及景观配置均呈现不同的特征,其中扶绥地区人为干扰最为强烈,德保地区的景观块数破碎化程度较为严重,而龙州地区的人为干扰程度最小,其森林景观最为聚集。(2)蔓延度指数、平均斑块分维指数、林地面积、林地斑块大小、裸岩面积和裸岩面积比重等景观指数与黑叶猴种群数量有显著正向关系,Shannon多样性指数则是显著负向关系;而耕地面积、耕地...     

Urbanisation versus agriculture: a comparison of local genetic diversity and gene flow between wood mouse Apodemus sylvaticus populations in human‐modified landscapes

Urbanisation and agriculture dramatically modify the landscapes available for use by wildlife, affecting key aspects of their ecology such as survival, foraging, predation, competition and reproductive success. Relatively little is known about the effects of urbanisation and agriculture on the genetic structure, gene flow and genetic diversity of wild species. Here, landscape genetic techniques were applied to compare local genetic diversity and gene flow between wood mouse populations in urban and arable landscapes. Using nine microsatellite markers, individuals were genotyped from six arable and seven urban sample sites. Inter‐population genetic differentiation was significantly greater in urban than arable habitat, while allele richness, private allele richness and heterozygosity were higher for arable sample sites, with varying degrees of significance. These suggest that urban habitat was sufficiently fragmented to limit gene flow. To test the effect of landscape features on gene flow, several cost‐distance measures were generated. Overland distance and Euclidean distance correlated best with inter‐population genetic differentiation in arable habitat, whereas distances that accommodated differences in habitat quality better explained differentiation in urban habitat. There was no evidence that margins adjacent to roads, rivers or railways facilitated gene flow. Together, the results indicate that urban landscapes expose wood mice to greater fragmentation in habitat quality than arable areas, leading to greater population isolation that is not mitigated by the presence of dispersal corridors.     

Assessing multi-taxa sensitivity to the human footprint,habitat fragmentation and loss by exploring alternative scenarios of dispersal ability and population size: a simulation approach

Quantifying the effects of landscape change on population connectivity is compounded by uncertainties about population size and distribution and a limited understanding of dispersal ability for most species. In addition, the effects of anthropogenic landscape change and sensitivity to regional climatic conditions interact to strongly affect habitat fragmentation and loss. To further develop conservation theory and to understand the interplay between all of these factors, we simulated habitat fragmentation and loss across the Western United States for several hypothetical species associated with four biome types, and a range of habitat requirements and dispersal abilities. We found dispersal ability and population size of the focal species to be equally sensitive to habitat extent, while dispersal ability is more sensitive to habitat fragmentation. There were also strong critical threshold effects where habitat connectivity decreased disproportionately to decreases in life-history traits making these species near these thresholds more sensitive to changes in habitat loss and fragmentation. Overall, grassland and forest associated species are also most at risk from habitat loss and fragmentation driven by human related land-use. These two largest biome types were most sensitive at large contiguous patch sizes which is often considered most important for metapopulation viability and biodiversity conservation. Hypothetical simulation studies such as this can be of great value to scientists in further conceptualizing and developing conservation theory, and evaluating spatially-explicit scenarios of habitat connectivity. Our results are available for download in a web-based interactive mapping prototype useful for accessing the results of this study.     

Parallel declines in species and genetic diversity in tropical forest fragments

The potential for parallel impacts of habitat change on multiple biodiversity levels has important conservation implications. We report on the first empirical test of the 'species-genetic diversity correlation' across co-distributed taxa with contrasting ecological traits in the context of habitat fragmentation. In a rainforest landscape undergoing conversion to oil palm, we show that depauperate species richness in fragments is mirrored by concomitant declines in population genetic diversity in the taxon predicted to be most susceptible to fragmentation. This association, not seen in the other species, relates to fragment area rather than isolation. While highlighting the over-simplification of extrapolating across taxa, we show that fragmentation presents a double jeopardy for some species. For these, conserving genetic diversity at levels of pristine forest could require sites 15-fold larger than those needed to safeguard species numbers. Importantly, however, each fragment contributes to regional species richness, with larger ones tending to contain more species.     

Landscape structure shapes habitat finding ability in a butterfly

Land-use intensification and habitat fragmentation is predicted to impact on the search strategies animals use to find habitat. We compared the habitat finding ability between populations of the speckled wood butterfly (Pararge aegeria L.) from landscapes that differ in degree of habitat fragmentation. Naïve butterflies reared under standardized laboratory conditions but originating from either fragmented agricultural landscapes or more continuous forested landscapes were released in the field, at fixed distances from a target habitat patch, and their flight paths were recorded. Butterflies originating from fragmented agricultural landscapes were better able to find a woodlot habitat from a distance compared to conspecifics from continuous forested landscapes. To manipulate the access to olfactory information, a subset of individuals from both landscape types were included in an antennae removal experiment. This confirmed the longer perceptual range for butterflies from agricultural landscapes and indicated the significance of both visual and olfactory information for orientation towards habitat. Our results are consistent with selection for increased perceptual range in fragmented landscapes to reduce dispersal costs. An increased perceptual range will alter the functional connectivity and thereby the chances for population persistence for the same level of structural connectivity in a fragmented landscape.     

Using population viability analysis,genomics, and habitat suitability to forecast future population patterns of Little Owl Athene noctua across Europe

The agricultural scene has changed over the past decades, resulting in a declining population trend in many species. It is therefore important to determine the factors that the individual species depend on in order to understand their decline. The landscape changes have also resulted in habitat fragmentation, turning once continuous populations into metapopulations. It is thus increasingly important to estimate both the number of individuals it takes to create a genetically viable population and the population trend. Here, population viability analysis and habitat suitability modeling were used to estimate population viability and future prospects across Europe of the Little Owl Athene noctua, a widespread species associated with agricultural landscapes. The results show a high risk of population declines over the coming 100 years, especially toward the north of Europe, whereas populations toward the southeastern part of Europe have a greater probability of persistence. In order to be considered genetically viable, individual populations must count 1,000–30,000 individuals. As Little Owl populations of several countries count <30,000, and many isolated populations in northern Europe count <1,000 individuals, management actions resulting in exchange of individuals between populations or even countries are probably necessary to prevent losing <1% genetic diversity over a 100‐year period. At a continental scale, a habitat suitability analysis suggested Little Owl to be affected positively by increasing temperatures and urban areas, whereas an increased tree cover, an increasing annual rainfall, grassland, and sparsely vegetated areas affect the presence of the owl negatively. However, the low predictive power of the habitat suitability model suggests that habitat suitability might be better explained at a smaller scale.